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We examined differences in the physiology and life history between dimorphs of the common pistachio psyllid, Agonoscena pistaciae (Burckhardt and Lauterer) (Hemiptera: Aphalaridae), and how they differ in elicitating host plant production of key metabolites and volatile compounds involved in the recruitment of herbivores and natural enemies. Summer morphs had higher activities of glutathione S-transferase, carboxylesterase, acetylcholinesterase, and cytochrome P450 monooxygenase, superoxide dismutase, catalase, peroxidase, phenoloxidase, and a higher total protein content compared to winter morphs, whereas the latter had higher amounts of lipid, carbohydrate, and glycogen. Winter morphs were heavier, with a higher chitin content and longer preoviposition period, but greater fecundity and longevity than summer morphs. A lower LC50 to thiamethoxam for winter morphs resulted in higher mortality following exposure to the recommended rate of this insecticide in a greenhouse trial. Feeding by winter morphs elicited more strongly the release of volatile compounds known to be attractive to other herbivores, whereas feeding by summer morphs elicited more strongly the release of volatiles implicated in the attraction of natural enemies. Feeding by psyllids increased the concentrations of nitrogenous compounds, carbohydrates, vitamins, and amino acids in plants, the winter morph eliciting larger changes and more improved host plant quality. We conclude that winter morphs are more vulnerable targets for chemical control in early spring, whereas management of summer morphs could rely more on conservation biological control.

Aphid cornicles are abdominal appendages that secrete an array of volatile and nonvolatile compounds with diverse ecological functions. The emission of alarm pheromones yields altruistic benefits for clone-mates in the aphid colony, which is essentially a superorganism with a collective fate. Secreted droplets also contain unsaturated triglycerides, fast-drying adhesives that can be lethal when smeared on natural enemies but more often impede their foraging efficiency. The longest cornicles have evolved in aphids that feed in exposed locations and are likely used to scent-mark colony intruders. Reduced cornicles are associated with reliance on alternative defenses, such as the secretion of protective waxes or myrmecophily. Root-feeding and gall-forming lifestyles provide protected feeding sites and are associated with an absence of cornicles. In some eusocial gall-formers, soldier morphs become repositories of cornicle secretion used to defend the gall, either as menopausal apterae that defend dispersing alatae or as sterile first instars that dispatch predators with their stylets and use cornicle secretions as a construction material for gall repair. Collectively, the evidence is consistent with an adaptive radiation of derived cornicle functions molded by the ecological lifestyle of the aphid lineage.

Baculoviruses are highly evolved parasites that genetically reprogram the developing phenotype of their host insect to produce a vessel for virus replication and dispersal. Here we show that larvae of Helicoverpa armigera infected with HearNPV accumulate glucose in the midgut, which reduces food consumption and alters the dynamics of pathways governing metabolism and immunity. We used transcriptomics to demonstrate the role of the insulin signaling pathway in regulating the HearNPV infection process. Dietary restriction decreased mortality of infected larvae and reduced viral replication prior to death, whereas dietary supplementation with glucose produced the opposite effects. The expression of most tricarboxylic acid cycle (TCA) and energy metabolism-related genes was reduced in infected larvae, whereas the expression of immunity-, glycolysis- and insulin-related genes was enhanced. Treatment of infected larvae with insulin increased their survival, reduced viral replication, and inhibited climbing behavior compared to a control treatment with DMSO, whereas RNAi suppression of the insulin receptor gene produced the opposite effects. Inhibition of glycolysis with dichloroacetate (DCA) promoted viral replication and accelerated larval death, but inhibition of the TCA cycle with 2-deoxyglucose (2-DG) did not, although both diminished climbing behavior. This work demonstrates that successful baculovirus infections hinge on metabolic reprogramming of the host and concurrent suppression of immune responses in the larval midgut, with the insulin signaling pathway mediating a trade-off between glucose metabolism and virus resistance.

Aphids parasitized in later instars can give birth to several nymphs before their reproduction is curtailed by the developing parasitoid. We examined the life histories of Aphis fabae Scopoli born to mothers parasitized by Lysiphlebus fabarum Marshall, and their suitability as subsequent hosts, to test the ‘fecundity compensation’ hypothesis. Maternal parasitism negatively impacted life history parameters, resulting in reduced estimates of population increase (rm, R0, and λ), and increased generation time (GT) and doubling time (DT). These impacts were greater when the larva developing in the mother turned out to be female rather than male, and greater still when mothers were superparasitized. Maternal parasitism produced aphids with shorter hind tibia (HTL), at birth and at maturity, but their developmental time was unaffected. Although female L. fabarum readily accepted such aphids for oviposition, rates of mummification and wasp emergence were lower, and more so when the maternal parasitoid was female. The resulting parasitoids took longer to develop than progeny from control wasps, had shorter HTLs, lower egg loads, smaller eggs, and produced fewer mummies with lower rates of adult emergence, all differences that were more pronounced when the maternal parasitoid was female. The progeny of these wasps exhibited similar impairments to these biological parameters as their parents, demonstrating that the negative impacts of development in maternally parasitized hosts extended for at least two generations. Thus, our results do not support fecundity compensation, but suggest that any benefits of post-parasitism reproduction will be offset by reduced fitness in both aphid progeny and the parasitoids that develop in them.

We studied the effect of Trichogramma brassicae Bezdenko (Hymenoptera: Trichogrammatidae) parasitism of Tuta absoluta Meyrick (Lepidoptera: Gelechiidae) eggs on the foraging behavior of Nabis pseudoferus Remane (Hemiptera: Nabidae) feeding on T. absoluta eggs of different ages. Older parasitized eggs (48–72 h old) were rejected after probing with the probosis. A separate experiment showed that probing did not harm developing parasitoids. Bugs greatly reduced their feeding times on older parasitized eggs and spent more time grooming and probing eggs and leaves. Predators exhibited a type III functional response to 24 h-old eggs, but a type II response to older eggs, whether parasitized or not. 24 h-old eggs suffered higher attack rates, and required less handling time than older eggs. Although the predatory efficiency of N. pseudoferus was reduced in the presence of parasitized eggs, the partial refuge enjoyed by T. brassicae in older eggs may permit an additive impact on the pest, and should facilitate the joint augmentation of both species against T. absoluta.

Ladybirds (Coleoptera: Coccinellidae) provide services that are critical to food production, and they fulfill an ecological role as a food source for predators. The richness, abundance, and distribution of ladybirds, however, are compromised by many anthropogenic threats. Meanwhile, a lack of knowledge of the conservation status of most species and the factors driving their population dynamics hinders the development and implementation of conservation strategies for ladybirds. We conducted a review of the literature on the ecology, diversity, and conservation of ladybirds to identify their key ecological threats. Ladybird populations are most affected by climate factors, landscape composition, and biological invasions. We suggest mitigating actions for ladybird conservation and recovery. Short-term actions include citizen science programs and education, protective measures for habitat recovery and threatened species, prevention of the introduction of non-native species, and the maintenance and restoration of natural areas and landscape heterogeneity. Mid-term actions involve the analysis of data from monitoring programs and insect collections to disentangle the effect of different threats to ladybird populations, understand habitat use by taxa on which there is limited knowledge, and quantify temporal trends of abundance, diversity, and biomass along a management-intensity gradient. Long-term actions include the development of a worldwide monitoring program based on standardized sampling to fill data gaps, increase explanatory power, streamline analyses, and facilitate global collaborations.Las catarinas (Coleoptera: Coccinellidae) proporcionan servicios que son críticos para la producción de alimento, y juegan un papel ecológico como fuente de alimento para depredadores. Sin embargo, la riqueza, abundancia y distribución de catarinas están en peligro debido a muchas amenazas antropogénicas. La carencia de conocimiento sobre el estatus de conservación de la mayoría de las especies y los factores que inciden en su dinámica poblacional dificulta el desarrollo e implementación de estrategias de conservación para las catarinas. Realizamos una revisión de la literatura sobre la ecología, diversidad y conservación de catarinas para identificar sus amenazas ecológicas clave. Las poblaciones de catarinas fueron afectadas mayormente por factores climáticos, composición del paisaje e invasiones biológicas. Proponemos acciones de mitigación para la conservación y recuperación de catarinas. Acciones a corto plazo incluyen programas de ciencia y educación ciudadana, medidas de protección para la recuperación de hábitat y de especies amenazadas, prevención de la introducción de especies no nativas y el mantenimiento y restauración de áreas naturales y la heterogeneidad del paisaje. Acciones a mediano plazo implican el análisis de datos obtenidos de programas de monitoreo y colecciones de insectos para desenmarañar el efecto de las diferentes amenazas a las poblaciones de catarinas, comprender el uso del hábitat por taxa de los que se tiene conocimiento limitado y cuantifica las tendencias temporales de la abundancia, diversidad y biomasa a lo largo de un gradiente de intensidad de manejo. Acciones a largo plazo incluyen el desarrollo de un programa de monitoreo a nivel mundial basado en muestreos estandarizados para subsanar la falta de datos, incrementar el poder explicativo, optimizar los análisis y facilitar colaboraciones globales.随着世界范围内人类与野生动物的冲突不断升级, 对野生动物的宽容和接纳等概念变得越来越重要。然而, 当代保护研究表明, 人们对人类与野生动物积极互作的认识有限, 导致可能难以准确描述人类与动物的相遇。如果不解决这些局限性, 就会导致野生动物和景观管理计划的设计和实施存在不足, 以及对本土生态知识的否定。我们利用印度喀拉拉邦瓦亚纳德野生动物保护区森林中的阿迪瓦西族Kattunayakans部落的民族志证据, 来研究印度原住民对于人类与野生动物共存的观点。通过定性的实地研究(包括访谈和在森林样带调查), 我们发现Kattunayakans部落表现出对野生动物的宽容和接纳, 其特点是深入的共存, 包括三个中心思想:野生动物是有理性的交流者;野生动物是神、老师和平等的族群;野生动物是实行 “dharmam”的具有共同起源的亲戚。我们认为, 充分理解以上几点有助于将Kattunayakan部落的观点引入印度的森林管理, 并更广泛地解决人类与野生动物的冲突。.

The entomopathogenic fungus Beauveria bassiana and the egg parasitoid Trichogramma dendrolimi can both contribute to biological control of the global fruit borer Grapholita molesta. To derive insights for optimizing their combined application in augmentation programs, we assayed fungal pathogenicity to both G. molesta (eggs, larvae and adults) and adult parasitoids, and assessed wasp acquisition and transmission of fungal spores following their emergence from B. bassiana-treated host eggs.Concentrations of 10Our results indicate that combined applications of B. bassiana and T. dendrolimi can have complimentary impacts on G. molesta, the wasps compensating for low fungal pathogenicity to eggs and their progeny potentially aiding in subsequent disease transmission. Although foraging wasps tended to avoid infected eggs, negative interactions between the two agents might be further mitigated by timing B. bassiana applications from late instar larva to early moth stage, and T. dendrolimi releases several days later to coincide with peak oviposition periods. © 2022 Society of Chemical Industry.

The convergent lady beetle, Hippodamia convergens Guerin-Meneville, is a specialized predator of cereal aphids on the High Plains, completing its first generation each year in winter wheat, the resulting adults dispersing into summer crops and producing additional generations, contingent on the availability of aphids. In the present study, we tested the collective value of supplementary plant resources (sugars, pollen, and seedling wheat leaves), and small amounts of alternative prey, eggs of Ephestia kuehniella (Zeller) (Lepidoptera: Pyralidae), all provided together, for improving beetle life history and reproductive success even when suitable prey, Schizaphis graminum (Rondani) (Hemiptera: Aphididae), were provided ad libitum. Although a monotypic diet of S. graminum yielded slightly faster larval development and heavier adults than the omnivorous diet, preoviposition periods were extended and 21-d fecundities greatly reduced when this diet was continued through adult life, largely due to fewer oviposition days, although egg fertility was unaffected. The results highlight the critical importance of plant-derived resources even when suitable prey are not limiting. However, monotypic diet beetles that diapaused for 21 d in the presence of supplementary plant resources, plus moth eggs, achieved the same reproductive success as those reared on the omnivorous diet, with or without diapause, demonstrating that access to these resources post-emergence was sufficient to compensate for their absence during development. The diapause treatment itself had no impact on the fitness of beetles reared on the omnivorous diet, likely because neither its duration, nor the caloric restriction imposed, were sufficient to diminish reproductive effort.

Although the brown stink bug, Euschistus heros (F.) is susceptible to entomopathogenic fungi, only a single formulation is currently available on the Brazilian market for its control. The objective...

The nutritional status of host plants can have direct impacts on herbivore physiology and insect–plant interactions. We investigated the effect of micronutrients, including manganese, iron, zinc, and copper, on cucumber plant physiology, and on the biology and physiology of a strain of Aphis gossypii Glover selected over 12 generations to be resistant to pirimicarb. The micronutrient treatment increased the activity of superoxide dismutase, ascorbate peroxidase, guaiacol peroxidase, polyphenol oxidase, and phenylalanine ammonia-lyase in cucumber plants, and also increased levels of total phenolics, hydrogen peroxide, salicylic acid, and total chlorophyl, whereas malondialdehyde levels were unaffected. Pirimicarb-resistant cotton aphids that fed on micronutritient-amended cucumber plants expressed significantly decreased levels of acetylcholinesterase and detoxifying enzymes, specifically glutathione S-transferase, and carboxylesterase. Analysis of energy reserves in resistant A. gossypii fed on micronutritient-amended plants revealed decreases in the lipid and protein contents of aphids, whereas glycogen and carbohydrate contents showed no response. Resistant cotton aphids fed on micronutritient-amended plants showed significantly reduced fecundity, longevity, and reproductive periods, and a 1.7-fold reduction in pirimicarb LC50 compared with those fed on control plants. We conclude that micronutrient amendment negatively impacts the biological performance of insecticide-resistant cotton aphids, and diminishes their resistance to pirimicarb. Both direct effects on plant health, such as enhanced inducible defenses, and indirect effects on aphid fitness, such as reduced biological performance and detoxification abilities, were implicated. Therefore, optimization of micronutrient amendments could be a useful complement to other tactics for managing insecticide-resistant A. gossypii on cucumbers, and warrants exploration in other contexts.

Body size is a trait with many potential impacts on fitness. Adult body size can affect the strength of condition-dependent parental effects that determine offspring phenotypes, with potentially important transgenerational consequences. In a preliminary experiment, larval food deprivation (30 min daily access) created Harmonia axyridis Pallas (Coleoptera: Coccinellidae) females that weighed ad libitum food (eggs of Ephestia kuehniella Zeller). Although only 1/3 of larvae survived to adulthood in the 30 min treatment, adult pairs produced eggs that were not significantly different in size from those of pairs fed ad libitum as larvae. Less extreme larval food deprivation (4 h daily access) was used to create a cohort of H. axyridis that weighed ad libitum food. Small couples had lower 20-day fecundities and reduced egg fertility relative to large couples. Both egg and pupal periods were shortest when both parents were small, and longest when both parents were large, with reciprocal crosses intermediate. There were no consistent effects of parental body size on larval development time, but the progeny of small females mated to large males pupated later than other treatments. Progeny of large pairs had the heaviest adult weights at emergence, and progeny of small pairs, the lightest, with the progeny of reciprocal crosses intermediate. Small females produced the lightest female offspring, whereas small males sired the lightest male offspring, suggesting stronger responses to epigenetic signals from parents of the same sex. These results indicate that H. axyridis cohorts maturing with abundant food will produce progeny with larger potential body size and fitness, whereas those experiencing food limitation will confer size and fitness limitations to the subsequent generation, with potentially important implications for short-term population dynamics.

Oenopia conglobata (Linnaeus) and Menochilus sexmaculatus Fabricius (Coleoptera: Coccinellidae) are important predators of the pistachio psyllid, Agonoscena pistaciae Burckhardt and Lauterer (Hemiptera: Psyllidae), a key pest of pistachio. The abundance of M. sexmaculatus has recently increased in pistachio orchards, while that of O. conglobata has declined. We designed laboratory experiments to (1) evaluate the potential for intraguild predation (IGP) between these two species, (2) detect any possible asymmetries in IGP interactions, and (3) characterize the sensitivity of IGP to varying densities of two extraguild (EG) prey, A. pistaciae and Aphis gossypii Glover (Hemiptera: Aphididae). We measured rates of IGP by fourth instar larvae and female adults (both starved for 12 h) on vulnerable immature life stages of the other species (eggs, first, and second instar larvae). IG and EG prey consumption were tallied after 12 h. All factors influenced rates of IGP, but species of IG predator, and life stage and density of EG prey, contributed the most variance. IGP decreased parabolically with increasing prey densities. IGP on eggs was higher than on first instars, which was higher than on second instars, suggesting palatability declined in later life stages. The larger M. sexmaculatus exhibited higher voracity than the smaller O. conglobata in both predation and IGP. Both species consumed more A. pistaciae than A. gossypii in all treatments, which may reflect either lower handling time or lower food value per prey.

Cotton bollworm, Helicoverpa armigera Hubner, is a cosmopolitan polyphagous pest of many crops. Habrobracon hebetor Say and Helicoverpa armigera nucleopolyhedrovirus (HearNPV) are two important biocontrol agents used to manage this pest, sometimes in combination. We evaluated the sublethal effects of HearNPV on H. hebetor life table parameters under laboratory conditions when its host (second instar H. armigera) was treated with HearNPV, and tested H. hebetor females for their ability to discriminate against inoculated hosts. Emergence of adults reared as solitary larvae was reduced by half on LC30-inoculated larvae compared to controls, but not on LC5 or LC15-inoculated hosts. Low concentrations (LC5, LC15, and LC30) of HearNPV had no effects on overall parasitoid developmental time, but longevity and lifetime fecundity was reduced for females emerging from hosts receiving the LC30 treatment. Net reproductive rate (R0), intrinsic rate of increase (rm), and finite rate of increase (λ) were all decreased in a concentration-dependent manner in the LC15 and LC30 treatments, as were female life expectancy, age-specific survivorship (lx), and age-specific fecundity (mx), whereas population doubling time (DT) increased. Parasitoids did not discriminate against LC15-inoculated larvae in choice or no-choice tests, but parasitized more LC50-inoculated hosts than controls in the choice test, with no significant differences in total numbers of eggs laid in either case. Although parasitoids suffered some loss of fitness in HearNPV-inoculated hosts under these laboratory conditions, these agents still appear compatible for joint application against H. armigera under field conditions, provided parasitoid releases are made 2 d after NPV application.

Invasive airborne insects pose major challenges in natural resource and agriculture management, as they can rapidly spread over large distances and cross physical boundaries. Field monitoring and local management are important tools to prevent and control infestations but require additional coordination to be operative region-wide. Computational modeling techniques have been effective in simulating local population dynamics and in capturing spread of invasive species on a regional scale. We use an integrated ecological model to simulate local and regional infestation dynamics of sugarcane aphids, Melanaphis sacchari (Zehntner) (Hemiptera: Aphididae), on sorghum, Sorghum bicolor (L.) Moench (family Poaceae), in the southern to central Great Plains of the United States. Local dynamics of aphid populations on sorghum are simulated by a spatially explicit, individual-based model, whereas regional aphid migration is simulated by an atmospheric model that computes inert air particle (aphid) transport, dispersion, and deposition. Simulation results indicate timing of initial infestations in the south affects spatiotemporal patterns of infestation throughout the region. Probability of local infestations is a function of both percentage of land occupied by growing sorghum and prevailing winds. Thus, due to availability of sorghum, relatively later dates of initial infestation in the south will probably lead to infestations farther north that become established more quickly following the first appearance of aphids in the south. The model we present, in coordination with field monitoring schemes, could be applied as a forecasting tool in region-wide pest management systems.

High temperatures will occur more frequently with global warming, with potential impacts on the efficacy of biological control agents. Heat shock proteins (HSPs) are induced by high temperature, but their possible roles in pest resistance to entomopathogens remain unexplored. We investigated the effects of high temperature (35 °C) on Helicoverpa armigera resistance to H. armigera nucleopolyhedrovirus (HearNPV) and the putative roles of HSPs in this process.Even short periods (24 h) of high temperature (35 °C) reduced mortality in HearNPV-infected H. armigera larvae. Sustained 35 °C exposure significantly shortened developmental time, and increased fresh weight and locomotor activity in infected larvae. Moreover, high temperature inhibited virus replication and thickened the epidermis of H. armigera, resulting in reduced spread of infection from cadavers. Real-time polymerase chain reaction (PCR) analysis showed that expression of 11 HSP genes was altered by the 35 °C treatment, and that mostly small heat shock protein (sHSP) genes were up-regulated, the same sHSPs were induced when larvae were infected with HearNPV. Finally, RNA interference (RNAi) suppression of these sHSPs showed that only Hsp24.91 and Hsp21.8 diminished H. armigera defensive responses to HearNPV infection.Even short periods of exposure to high temperature can significantly reduce susceptibility of H. armigera larvae to HearNPV by stimulating the production of sHSPs which enhance immune responses, with important implications for the use of entomopathogens as biological control agents under global warming scenarios. © 2022 Society of Chemical Industry.

Baculoviruses can induce climbing behaviour in their caterpillar hosts to ensure they die at elevated positions to enhance virus transmission, providing an excellent model to study parasitic manipulation of host behaviour. Here, we demonstrate that climbing behaviour occurred mostly during daylight hours, and that the height at death of Helicoverpa armigera single nucleopolyhedrovirus (HearNPV)-infected larvae increases with the height of the light source. Phototaxic and electroretinogram (ERG) responses were enhanced after HearNPV-infection in host larvae, and ablation of stemmata in infected larvae prevented both phototaxis and climbing behaviour. Through transcriptome and quantitative PCR, we confirmed that two opsin genes (a blue light-sensitive gene, HaBL; and a long wave-sensitive gene, HaLW) as well as the TRPL (transient receptor potential-like channel protein) gene, all integral to the host's visual perception pathway, were significantly upregulated after HearNPV infection. Knockout of HaBL, HaLW, or TRPL genes using the CRISPR/Cas9 system resulted in significantly reduced ERG responses, phototaxis, and climbing behaviour in HearNPV-infected larvae. These results reveal that HearNPV alters the expression of specific genes to hijack host visual perception at fundamental levels—photoreception and phototransduction—in order to induce climbing behaviour in host larvae.

As an intermediate molecule in the Insulin/Insulin-like growth factor signalling pathway (IIS), the insulin receptor (IR) plays vital roles linking nutritional signals to the downstream regulation of metabolic homeostasis, development, metamorphosis, reproduction and stress responses. In the present study, we describe the molecular characteristics of IR in the cosmopolitan fruit boring pest, Grapholita molesta, and its predicted posttranscription regulator miR-982490, and elucidate its regulatory roles in glucolipid homeostasis and metamorphosis. Phylogenetic and domain analyses indicate that lepidopteran IRs normally cluster within families, and that four main domains are conserved in GmIR and those of other Lepidoptera. Bio-informatic prediction, synchronic expression profile evaluation and dual luciferase reporter assays indicated negative regulation of GmIR by miR-982490. Injection of miR-982490 agomir into fifth instar larvae yielded effects similar to dsGmIR injection, resulting in enhanced levels of trehalose and triglyceride in haemolymph, and reduced pupation success and pupal weight, both of which could be rescued by co-injection of dsGmIR and miR-982490 antagomir. We infer that GmIR regulates glucolipid homeostasis and affects G. molesta metamorphosis via interactions with its posttranscriptional regulator miR-982490. This study expands our understanding of the regulatory network of IIS in insect nutritional homeostasis and development.

Adult aphid predators disperse across the landscape seasonally in search of prey aggregations that are patchily distributed and temporally variable. However, flight is energetically costly and consumes resources that could be invested in reproduction. Hippodamia convergens is an important aphid predator in North American cereal crops and other agricultural systems. Consumption of floral resources can enhance adult survival during periods of low prey availability and may improve reproductive success. We tested how an omnivorous adult diet containing floral resources (diluted honey and pulverized bee pollen) interacts with body size to influence reproduction and flight behavior compared to a prey-only diet. Two sizes of beetles were produced by controlling larval access to food—3 h daily access produced small beetles; ad libitum access produced large beetles with faster development. Reproductive performance was tracked for 18 days, and female flight activity was assayed via 3 h bouts of tethered flight. Diet composition and body size interacted to influence preoviposition period, with large females in prey-only treatments delaying oviposition the longest. The omnivorous adult diet improved 18-day fecundity relative to a prey-only diet, but egg fertility was unaffected. Adult size affected oviposition pattern, with small beetles laying smaller, but more numerous, clutches. Females flew up to 7 km in 6 h, but neither body size nor adult diet influenced flight distance, suggesting that all diet treatments generated energy reserves sufficient to power flights of short duration. However, pre-reproductive females flew > 60% further than they did post-reproduction, likely due to the energetic costs of oviposition. Thus, access to pollen and nectar increased reproductive success and altered oviposition patterns in H. convergens, indicating the importance of floral resources in the agricultural landscape to conservation of this predator and its biological control services.

The sugarcane aphid, Melanaphis sacchari (Zehntner) (Hemiptera: Aphididae) (SCA), has become a major pest of grain sorghum since its appearance in the USA. Several grain sorghum parental lines are moderately resistant to the SCA. However, the molecular and genetic mechanisms underlying this resistance are poorly understood, which has constrained breeding for improved resistance. RNA-Seq was used to conduct transcriptomics analysis on a moderately resistant genotype (TAM428) and a susceptible genotype (Tx2737) to elucidate the molecular mechanisms underlying resistance. Differential expression analysis revealed differences in transcriptomic profile between the two genotypes at multiple time points after infestation by SCA. Six gene clusters had differential expression during SCA infestation. Gene ontology enrichment and cluster analysis of genes differentially expressed after SCA infestation revealed consistent upregulation of genes controlling protein and lipid binding, cellular catabolic processes, transcription initiation, and autophagy in the resistant genotype. Genes regulating responses to external stimuli and stress, cell communication, and transferase activities, were all upregulated in later stages of infestation. On the other hand, expression of genes controlling cell cycle and nuclear division were reduced after SCA infestation in the resistant genotype. These results indicate that different classes of genes, including stress response genes and transcription factors, are responsible for countering the physiological effects of SCA infestation in resistant sorghum plants.

The egg parasitoid Trissolcus vassilievi (Mayr) is a key natural enemy of sunn pest, Eurygaster integriceps Puton (Hemiptera: Scutelleridae). This laboratory study examined how the aggregation of parasitoids varied in response to host density and numbers of foraging conspecifics. Five host densities (1, 2, 4, 8, and 15 clutches of E. integriceps eggs), were offered simultaneously to 1, 2, 4, 8, or 16 female wasps in a standardized arena (30 cm diameter) and patch residence times were recorded. Patch residence time was strongly correlated with parasitism, and increased as the number of host clutches increased up to eight, but declined when more clutches were offered. Wasps displayed low mobility and tended to remain in the patch initially encountered, even when it contained few egg masses. At higher wasp densities, patches were occupied more quickly, host exploitation began sooner, and per capita handling time was reduced. However, total patch residence times were similar across all densities. Thus, females responded to conspecifics with increased movement, which increased dispersal and resulted in a more homogeneous distribution of wasps among patches. Pseudo-interference, resulting from wasps remaining in some patches and neglecting others, had greater impact on final levels of host exploitation than did actual interference (patch abandonment following conspecific encounters). These phenomena resulted in decreased parasitoid searching rates and a 2-fold increase in host survival at higher wasp densities.

The chickpea pod borer, Helicoverpa armigera Hubner (Lepidoptera: Noctuidae), is an important pest of many legume crops, including chickpea, Cicer arietinum L. (Fabaceae). We field‐tested the efficacy of two pathogens, Helicoverpa armigera nucleopolyhedrosis virus (HaNPV) and Bacillus thuringiensis Berliner subsp. kurstaki (Btk), both alone and in combination with the parasitoid Habrobracon hebetor Say (Hh) (Hymenoptera: Braconidae), for control of H. armigera larvae in chickpea fields. A randomized complete block design was used at two sites, so that wasp release treatments at one site would not contaminate pathogen‐only treatments at the other. Pathogens were applied when most pest larvae were in the second instar, and wasp releases were made 2 days later. Number of H. armigera larvae per plant, number of damaged pods, yield (kg ha⁻¹), treatment cost (US$ ha⁻¹), and benefit:cost ratio were all determined. Treatments separated significantly at 14 days after treatment, with the HaNPV+Hh and Btk+Hh treatments providing the best control, suggesting a synergistic impact of the wasp and the pathogens when applied together. These treatments also produced the fewest damaged pods, the highest yields, and the greatest yield increments in comparison to untreated controls. However, costs were also highest for these combined treatments, which reduced their net benefit in comparison to use of Hh or Btk alone, although more efficient mass production of the pathogens might make combined applications feasible. We conclude that both HaNPV and Btk are suitable for integration with releases of H. hebetor against H. armigera in chickpea fields, provided parasitoid releases are delayed until 2 days after pathogen application.

Cotesia flavipes Cameron (Hymenoptera: Braconidae) is often augmented against Diatraea spp. (Lepidoptera: Crambidae) stem borers in the Americas. Diatraea indigenella Dyar and Heinrich, an endemic pest of sugarcane in southern Colombia, represents a novel host for C. flavipes. We examined the development of C. flavipes on D. indigenella and D. saccharalis, a familiar host, when larvae were stung by one, two or three wasps. Host species had no effect on parasitism success, immature parasitoid mortality, parasitoid developmental time, adult emergence, or female size. Superparasitized D. indigenella hosts yielded more cocoons, more adults, and more female wasps than those parasitized with a single sting, or from D. saccharalis hosts. However, wasp survival and longevity declined when host larvae were stung three times. Thus, D. indigenella was a more suitable host for C. flavipes when superparasitized, and appears suitable for augmentation against this emergent pest and other Diatraea species attacking sugarcane.

The nutritional suitability of prey is an important determinant of predator fitness that can have a cascade of effects on life history. We manipulated the quality of cucumber plants by fertilizing them with three rates of nitrogen, 110, 160 and 210 ppm N, representing low, medium and high levels of N fertilization, respectively. Colonies of Aphis gossypii Glover were reared on these plants and used as prey for Hippodamia variegata (Goeze) in experiments that measured the predator's development, reproduction, and predation rate. Biological parameters were calculated and an age-stage, two-sex life table was constructed. Larval development was extended and aphid consumption increased at the low level of N fertilization, likely due to the reduced N content of the aphids. However, female longevity was greatest in this treatment, likely because of a reduced reproductive rate, and there was no compensatory consumption of aphids by adults, as observed in larval stages, possibly due to carry-over effects of poor larval nutrition. Despite feeding on aphids with the highest N content, the high N treatment did not yield the highest intrinsic rate of increase (r). Rather, the medium N treatment (160 ppm N) yielded the highest aphid consumption rate and r value, and the highest efficiency of conversion of aphids into eggs.

Lydella minense and Billaea claripalpis (Diptera: Tachinidae) are important parasitoids used in augmentation biological control of sugarcane stem borers, Diatraea spp. (Lepidoptera: Crambidae), the primary pests of sugarcane in Colombia. Following the recent discovery of two new Diatraea species in the Cauca River Valley (CRV), D. tabernella and D. busckella, stalk-boring damage to sugarcane has increased and rates of larval parasitism have declined. The present study assessed the suitability of all four stalk-boring Diatraea species now present in the CRV, D. tabernella, D. busckella, D. indigenella and D. saccharalis, as hosts for the two parasitoids. Following manual inoculation of larvae with maggots of L. minense and B. claripalpis, the development, immature survival, sex ratio and longevity of resulting flies were recorded. Both parasitoid species parasitized a lower percentage of D. tabernella and D. busckella larvae compared to larvae of D. saccharalis or D. indigenella. Although host larvae were more likely to yield two puparia of L. minense than of B. claripalpis, the likelihood of a second puparium was reduced when L. minense parasitized D. tabernella and D. busckella. These results suggest these two Diatraea species have resistance to parasitism by the tachinids, which might explain why outbreaks of these pests have occurred in the regions of the CRV where commercial releases of the parasitoids are relied upon as the primary pest management tactic. Although breeding programs may eventually provide flies with improved ability to parasitize these novel host species, additional work is warranted to explore alternative pest management tactics for these stem borers.

Sorghum (Sorghum bicolor (L.) Moench) is an important summer grain crop in the U.S. Southern Great Plains because it is one of the few crops that consistently produces acceptable yields in the harsh summer weather that characterizes the region. Damaging infestations of the sugarcane aphid, Melanaphis sacchari (Zehntner), occur commonly in sorghum throughout Texas and Oklahoma, two of the leading sorghum producing states. Grain sorghum fields were sampled at two locations in Texas and three locations in Oklahoma. Fields were sampled approximately weekly by examining two leaves on each of 54 plants and counting all sugarcane aphids and aphid natural enemies on each leaf. Sampling began at an early growth stage and continued until the crop either matured or was treated with insecticide to suppress the sugarcane aphid infestation. For a total of 123 fields in 2017 and 2018, aphids, Aphelinus nigritus Howard mummies, adult coccinellids, larval coccinellids, larval lacewings, and larval dipterans were counted, and relative density estimates were determined. When natural enemy and sugarcane aphid count data were aggregated at the scale of geographic locations and multiple years, there was no evidence for a numerical response by natural enemies to sugarcane aphid density. When fields were compared within locations, a numerical response was consistently observed. A natural enemy importance index was developed that incorporated cumulative degree days of first occurrence of a natural enemy taxa in a field, the average density of the natural enemy in a field, and natural enemy voracity. Factorial analyses of variance indicated that cumulative degree days at first occurrence and average relative density differed significantly among natural enemy taxa and locations, as did natural enemy importance. Averaged across locations larval coccinellids had the largest importance index, I = 1.47, and A. nigritus had the smallest, I = 0.61. Among locations, the Texas Coastal Plains had the largest importance index, I = 1.27 whereas the Oklahoma Panhandle had the smallest, I = 1.02. Results suggest that differences occur in the biological control contributions of various natural enemies and in biological control efficacy among locations.

Background Oriental fruit moth, Grapholita molesta (Busck) (Lepidoptera: Tortricidae) is a cosmopolitan pests of fruit trees in temperate regions. Control of G. molesta is challenging because larvae feed in concealed locations and have evolved resistance to many pesticides. We compared three commercially available species of Trichogramma for efficacy against G. molesta in the laboratory assays and tested releases of the promising species in a pear orchard. Results Laboratory assays indicated that Trichogramma dendrolimi parasitized G. molesta at the highest rate. Parasitoids took longer to oviposit in older host eggs, and fewer eggs were parasitized when they were more than 3 days old. Field tests produced ca 60% cumulative parasitism of sentinel G. molesta eggs with one release of Trichogramma dendrolimi, with most parasitism occurring within 24 h. Female wasps dispersed up to 12 m from release points with a bias toward upwind movement. Conclusion We calculated that, for each generation of G. molesta, 900 000 wasps per hectare, in three releases 3 days apart, each distributed among release points 10 m apart, would reduce fruit damage by half in an orchard where 50% of fruit would otherwise be damaged. Although augmentation of Trichogramma dendrolimi is a viable tactic for reducing G. molesta populations and fruit damage, it will require integration with other compatible control tactics in order to provide commercially acceptable levels of control in orchards experiencing significant pest pressure. © 2021 Society of Chemical Industry.

Body size is a trait with many potential impacts on fitness. Adult body size can affect the strength of condition-dependent parental effects that determine offspring phenotypes, with potentially important transgenerational consequences. In a preliminary experiment, larval food deprivation (30 min daily access) created Harmonia axyridis Pallas (Coleoptera: Coccinellidae) females that weighed50% of controls reared on ad libitum food (eggs of Ephestia kuehniella Zeller). Although only 1/3 of larvae survived to adulthood in the 30 min treatment, adult pairs produced eggs that were not significantly different in size from those of pairs fed ad libitum as larvae. Less extreme larval food deprivation (4 h daily access) was used to create a cohort of H. axyridis that weighed60% of controls reared on ad libitum food. Small couples had lower 20-day fecundities and reduced egg fertility relative to large couples. Both egg and pupal periods were shortest when both parents were small, and longest when both parents were large, with reciprocal crosses intermediate. There were no consistent effects of parental body size on larval development time, but the progeny of small females mated to large males pupated later than other treatments. Progeny of large pairs had the heaviest adult weights at emergence, and progeny of small pairs, the lightest, with the progeny of reciprocal crosses intermediate. Small females produced the lightest female offspring, whereas small males sired the lightest male offspring, suggesting stronger responses to epigenetic signals from parents of the same sex. These results indicate that H. axyridis cohorts maturing with abundant food will produce progeny with larger potential body size and fitness, whereas those experiencing food limitation will confer size and fitness limitations to the subsequent generation, with potentially important implications for short-term population dynamics.

Bacillus thuringiensis Berliner subsp. kurstaki (Btk) and Habrobracon hebetor Say are both biological control agents of Helicoverpa armigera Hubner. The present study evaluated their compatibility for combined application against this pest by examining the acceptability of Btk-inoculated hosts for H. hebetor females and testing for negative life-history impacts on developing progeny. Second-instar H. armigera larvae fed for 72 h on potted chickpea plants treated with three concentrations of Btk (LC15, LC35, and LC70) and were then used in bioassays of parasitoid development and parasitism behavior. Survival of parasitoids was significantly reduced, and immature development prolonged, on hosts fed chickpea plants treated with LC35 and LC70 Btk, but not on plants treated with LC15 Btk. Parasitoids failed to discriminate against hosts treated with LC15 or LC35 Btk in choice tests, but attacked fewer hosts treated with LC70 Btk, paralyzing and parasitizing more healthy hosts, and laying more eggs on them. In contrast, a no-choice test revealed that more hosts treated with LC35 and LC70 Btk were paralyzed compared with control or LC15-treated hosts, but the numbers of hosts parasitized and eggs laid did not vary among Btk treatments. Thus, females required an experience with healthy hosts, as they had in the choice test, to discriminate against diseased ones. We conclude that H. hebetor and Btk are compatible for joint application against H. armigera, which could potentially improve biological control of this pest.

A standardized laboratory procedure tested the limb regeneration abilities of 18 populations (16 species) of lady beetles which were then scored, relative to unoperated controls, based on survival, the proportion regenerating the limb completely or partially, and the magnitude of developmental costs (delayed development, reduced body size) associated with limb ablation. Newly molted fourth instar larvae each had a single foreleg amputated at the base of the femur. All species except Propylea dissecta (Mulsant) showed some complete limb regeneration, with limb regeneration index (LRI) scores ranging from 0.025 to 0.905 out of a possible 1.00 (mean = 0.598). Eriopis connexa Germar, an aphidophagous neotropical species, scored the highest. Widely distributed species that dominate agricultural habitats all scored above 0.75, and the only herbivore, Henosepilachna argus (Geoffroy), scored second from lowest. Prolonged pupal development was the most common cost, occurring in nine species, and correlating negatively with regeneration. Taxonomic distance between species correlated with regeneration, but explained

Salicylic acid (SA) is a signaling molecule that can induce plant resistance to certain herbivores. Although the role of jasmonic acid in mediating mite-tomato plant interactions has been well studied, the role of salicylic acid has not. This study examined how the application of exogenous SA, via its effects on tomato plant physiology, alters the activity of mite digestive enzymes, mite energy reserves, and mite susceptibility to spirodiclofen. Enzymatic activity—including superoxide dismutase, ascorbate peroxidase, guaiacol peroxidase, polyphenol oxidase, and phenylalanine ammonia-lyase—along with contents of total phenolic, hydrogen peroxide, and total chlorophyll significantly increased in plants 24 h after treatment with 2 mM of SA. In contrast, catalase activity significantly decreased in treated plants, and malondialdehyde content was unaffected. Mites fed on tomato plants treated with SA had significantly lower glutathione S-transferase, esterase, α-amylase, and aminopeptidase activities than those fed on control plants. Energy reserve analyses demonstrated a significant decrease in contents of lipid, protein, and glycogen in mites fed on SA-treated plants, whereas carbohydrate content significantly increased. The LC50 of spirodiclofen was decreased 1.8-fold for Tetranychus urticae fed on SA-treated tomato plants compared to controls. Treatment of adult mites with 2 mM SA on leaf discs did not cause any direct mortality after 24 h. Finally, a greenhouse bioassay confirmed that spider mite mortality following exposure to spirodiclofen was significantly higher on SA plants than on control plants. Mortality of mites exposed to half of the recommended rate of spirodiclofen was similar to those exposed to the recommended rate when they were held on treated plants. These results have valuable implications for T. urticae management programs in tomato production.

Grapholita molesta, the oriental fruit moth, is a serious global pest of many Rosaceae fruit trees. Gut microorganisms play important roles in host nutrition, digestion, detoxification, and resistance to pathogens. However, there are few studies on the microbiota of G. molesta, particularly during metamorphosis. Here, the diversity of gut microbiota across the holometabolous life cycle of G. molesta was investigated comprehensively by Illumina high-throughput sequencing technology. The results showed that the microbiota associated with eggs had a high number of operational taxonomic units (OTUs). OTU and species richness in early-instar larvae (first and second instars) were significantly higher than those in late-instar larvae (third to fifth instars). Species richness increased again in male pupae and adults, apparently during the process of metamorphosis, compared to late-instar larvae. Proteobacteria and Firmicutes were the dominant phyla in the gut and underwent notable changes during metamorphosis. At the genus level, gut microbial community shifts from Gluconobacter and Pantoea in early-instar larvae to Enterococcus and Enterobacter in late-instar larvae and to Serratia in pupae were apparent, in concert with host developmental changes. Principal coordinate analysis (PCoA) and linear discriminant analysis effect size (LEfSe) analyses confirmed the differences in the structure of gut microbiota across different developmental stages. In addition, sex-dependent bacterial community differences were observed. Microbial interaction network analysis showed different correlations among intestinal microbes at each developmental stage of G. molesta, which may result from the different abundance and diversity of gut microbiota at different life stages. Phylogenetic Investigation of Communities by Reconstruction of Unobserved States (PICRUSt) analysis indicated that most functional prediction categories of gut microbiota were related to membrane transport, carbohydrate and amino acid metabolism, and DNA replication and repair. Bacteria isolated by conventional culture-dependent methods belonged to Proteobacteria, Firmicutes, and Actinobacteria, which was consistent with high-throughput sequencing results. In conclusion, exploration of gut bacterial community composition in the gut of G. molesta should shed light into deeper understanding about the intricate associations between microbiota and host and might provide clues to the development of novel pest management strategies against fruit borers.

Conservation biological control (CBC) makes valuable contributions to productivity in High Plains agriculture. Despite some notable exceptions, CBC holds a wide range of potentially damaging pests under natural control, largely due to the actions of beneficial arthropod species that have endured more than a century of forced adaptation to an increasingly industrialized agricultural landscape. 'Evolutionary rescue' is a natural selection process whereby species evolve adaptations that enable them to survive environmental conditions that would have been lethal to their ancestors. This process has structured both pest populations and the guilds of natural enemies that help suppress them in our highly disturbed agroecosystems. Rescued beneficial species, by virtue of their abundance and successful evolutionary history, including some either accidentally or intentionally introduced, are those with the greatest potential for future adaptation in the face of changing cultural practices and the emergence of novel pests. I review some existing impediments to CBC (large-scale, synchronous monocultures, inadequate plant and insect diversity in the landscape, migratory pests, vulnerable crop cultivars that rescue pests, rather than their enemies, and insecticidal seed treatments). Compounding these problems are commercial interests and agricultural philosophies that tend to incentivize yield-maximization and short-term profits over long term farm sustainability and profitability. However, certain agronomic practices implemented recently have likely benefited CBC. These include improved host plant resistance in crop cultivars, the advent of Bt-protected crops (which have reduced pesticide applications), the adoption of no-till and reduced tillage practices, more advanced irrigation technologies, and the development of more selective pesticides that are safer for beneficial arthropods. Looking forward, two current trends may improve the efficacy of CBC in row crops; the increased adoption of cover crops as an alternative to chemfallow periods, and increasing public concern for preservation of pollinator habitat on public lands, which will also benefit predators and parasitoids.

The American serpentine leaf miner, Liriomyza trifolii (Burgess) (Diptera: Agromyzidae), is a perennial pest of leguminous crops in the Mediterranean region. A life table was constructed for L. trifolii infesting broad bean, Vicia faba L., in northern Egypt. Key factor analysis was used to rank sources of immature mortality over two seasons. Leaf miners had three successive generations, and a partial fourth, in each year, with peak abundance in March-April. Less than 15 and 22% of L. trifolii survived to adult in seasons one and two, respectively. The largest contributor of immature leaf miner mortality in both seasons was unknown (41.2 and 39.1% of total mortality, respectively), and likely comprised a combination of abiotic factors, parasitoid-inflicted mortality (host-feeding), and predation. Parasitism was second, contributing 36.2 and 35.6% of total mortality in the two seasons, respectively, primarily due to larval parasitism by Diglyphus isaea (Walker) (Hymenoptera: Eulophidae), and low levels of larval-pupal parasitism by Opius pallipes Wesmael (Hymenoptera: Braconidae) and Halticoptera circulus (Walker) (Hymenoptera: Pteromalidae). Residual mortality resulted from malformed pupae or failed adult emergence. Key factor analysis revealed malformation to be the major cause of pupal mortality. Sequential regression confirmed that unknown mortality and D. isaea were the top stage-specific factors, both acting on larvae. Parasitoid abundance tracked host abundance across generations, but density dependence was not observed for any mortality factor, and the magnitudes of regression slopes were small. The results indicate the potential importance of conservation biological control in management of L. trifolii, given that naturally occurring parasitoids and other biotic/abiotic factors exert significant mortality on immature leaf miners.

1. For this study a cohort of melanic Harmonia axyridis males homozygous for the spectabilis allele was produced. These were used to produce four kinds of twice‐mated females, comprising all four permutations of melanic and non‐melanic (succinic) males. A series of 12 larvae were then reared from the first and 10th clutches of each female to compare progeny developmental phenotypes. 2. There were no effects of mating treatment on overall female reproductive performance (preoviposition period, 20‐day fecundity, or egg fertility). 3. Age‐specific maternal effects were evident in progeny developmental phenotypes; larvae of 10th clutches developed more slowly, pupation was shorter, and adults emerged at heavier weights. 4. Paternal effects were superimposed on maternal effects and affected progeny independent of their paternity; melanic males induced slower larval development and slower pupation, but only in 10th clutches and only when they mated second. 5. There was a significant three‐way interaction between male mating treatment, clutch number and progeny phenotype, indicating that progeny developmental responses to (mixed) paternal effects varied depending on their own phenotype and the time elapsed since their mother's last mating. 6. Melanic males mating second obtained a P2 advantage over succinic males, which increased from first to 10th clutch, but the reverse was not true when succinic males mated second. Thus, polyandry in H. axyridis facilitates both genetic and epigenetic competition among males while simultaneously enabling the sharing of predominant paternal effects among the progeny of different fathers.

Gaeolaelaps aculeifer (Canestrini, 1883) is a soil-dwelling predatory mite with potential for use as a biological control agent of fungus gnats (Diptera: Sciaridae) in mushroom production. The life...

Augmentation biological control has successfully replaced a lot of insecticide use in 'closed system' agriculture (e.g., greenhouses). The profitable commercialization of biocontrol agents in greenhouses has created an incentive to expand markets for mass-reared beneficial insects into open agricultural systems, often without sufficient scientific justification. However, the semi-contained nature of greenhouse culture is often critical to the success of augmentation and can serve to mask potential pitfalls and intrinsic limitations of this approach in open systems. Factors contributing to greenhouse successes include the reduced biological diversity of contained agroecosystems, the prevention of agent dispersal, the ability to maintain environmental conditions within a range favorable for the agent, the exclusion of competitors and natural enemies of the agent that might otherwise diminish its efficacy, and the absence of alternative prey/hosts that could divert predation/parasitism from the target pest. There are also problems arising from collection of source material from locally adapted populations, and the inadvertent imposition of artificial selection in the course of laboratory rearing. Besides highlighting these pitfalls, this paper aims to encourage more consideration of conservation approaches prior to investment in augmentation programs which entice farmers into perpetual cycles of 'rear and release.' I argue that although augmentation can benefit agriculture whenever it replaces pesticide applications, it does not constitute an ecologically sustainable solution because it requires continued inputs, and it can distract research attention away from more sustainable objectives. Sustainable biological control is best achieved through modifications to cultural practices that increasingly 'naturalize' agroecosystems, thus facilitating the natural recruitment and persistence of beneficial arthropod fauna, combined with habitat management geared to increasing overall plant and arthropod diversity in the agroecosystem.

1. The fitness impacts of two levels of superparasitism were compared in a sexual and an asexual strain of Lysiphlebus fabarum (Marshall) developing in black bean aphids. 2. Asexual females were larger than sexuals, had higher eggs loads, and better adult emergence in two of three treatments, but parasitised fewer available aphids in 24 h (80% fewer when foraging alone). 3. Superparasitism resulted in wasps that were larger than those emerging from singly parasitised aphids, and asexual females had larger ovaries, without delayed development. In contrast, sexual wasps took about 2 days longer to develop in superparasitised hosts, and females had smaller ovaries with lower egg loads. 4. The growth of host aphids bearing asexual larvae was not reduced relative to healthy aphids, except when heavily superparasitised, whereas sexual larvae reduced aphid growth in all treatments. 5. Elimination of supernumerary larvae was virtually complete at 4.5 days after parasitism by sexual females, but no elimination occurred among asexual larvae. The evolution of diminished aggression in asexual larvae may be facilitated by greater genetic similarity, without necessarily leading to gregarious development, assuming the extended life of supernumeraries somehow improves survivor fitness. 6. Sons of virgin sexual females developed faster than sons of mated females, implicating parental effects that slowed development of the latter. 7. It is concluded that asexual females of this strain use superparasitism to improve host quality, and thus the fitness of their daughters, whereas females of the sexual strain prioritise offspring number.