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The psyllid Cacopsylla chinensis (Yang & Li) (Hemiptera: Psyllidae) is a serious pest of pears in China. To determine and contrast the fitness of the psyllid on two endemic cultivars of Pyrus bretschneideri (i.e., BHXS and BSL) and two introduced cultivars of Pyrus communis (i.e., CB and CRB), we analyzed data on the development, survival, and fecundity from C. chinensis individuals reared on the four cultivars. The age-stage, two-sex life table theory was used in order to enable the inclusion of males in the analysis as well as a means of identifying the variation in developmental durations among individuals. Results indicated that C. chinensis can successfully develop and reproduce on all four pear cultivars. However, based on the lower preadult survival rate, longer preadult duration, longer total preoviposition period (TPOP), and lower fecundity that occurred on both cultivars of P. communis, these two cultivars are less favorable hosts for C. chinensis than the P. bretschneideri cultivars. The lower intrinsic rate of increase (r), finite rate of increase (λ), and net reproduction rate (R0) on CB and CRB pears showed these two introduced cultivars are more resistant to C. chinensis than the endemic BHXS and BSL pears. These resistant cultivars would be appropriate candidates for managing C. chinensis. We used the bootstrap technique to estimate the uncertainty of the population parameters (r, λ, R0, etc.), while also demonstrating that it can be used for estimating the 0.025 and 0.975 percentile confidence intervals of the age of survival rate. Graphical Abstract Age-stage, two-sex life table can reveal the survial and stage differentiation for assessment of differences between treatments

Huanglongbing (HLB) is a bacterial infection of citrus trees transmitted by the Asian citrus psyllid Diaphorina citri . Mitigation of HLB has focused on spraying of insecticides to reduce the psyllid population and removal of trees when they first show symptoms of the disease. These interventions have been only marginally effective, because symptoms of HLB do not appear on leaves for months to years after initial infection. Limited knowledge about disease spread during the asymptomatic phase is exemplified by the heretofore unknown length of time from initial infection of newly developing cluster of young leaves, called flush, by adult psyllids until the flush become infectious. We present experimental evidence showing that young flush become infectious within 15 d after receiving an inoculum of Candidatus Liberibacter asiaticus (bacteria). Using this critical fact, we specify a microsimulation model of asymptomatic disease spread and intensity in a grove of citrus trees. We apply a range of psyllid introduction scenarios to show that entire groves can become infected with up to 12,000 psyllids per tree in less than 1 y, before most of the trees show any symptoms. We also show that intervention strategies that reduce the psyllid population by 75% during the flushing periods can delay infection of a full grove, and thereby reduce the amount of insecticide used throughout a year. This result implies that psyllid surveillance and control, using a variety of recently available technologies, should be used from the initial detection of invasion and throughout the asymptomatic period. Significance Huanglongbing (HLB) is a vector-transmitted bacterial infection of citrus trees that poses a major threat to the citrus industry in Florida, Texas, and California. Current control strategies that focus on the vector, the Asian citrus psyllid Diaphorina citri , are usually initiated when the trees become symptomatic, anywhere from 10 mo to several years after initial infection. We show, experimentally, that newly infected young leaves can become infectious within 10–15 d after receiving an inoculum of bacteria from an adult psyllid. We then show by microsimulation of the asymptomatic spread of HLB through a grove under different invasion scenarios and control strategies that reduction of up to 75% of adult psyllids and nymphs can enhance citrus production.

Wolbachia pipientis is a maternally inherited intracellular bacterium that infects a wide range of arthropods. Wolbachia can have a significant impact on host biology and development, often due to its effects on reproduction. We investigated Wolbachia-mediated effects in the Asian citrus psyllid, Diaphorina citri Kuwayama, which transmits Candidatus Liberibacter asiaticus (CLas), the causal agent of citrus greening disease. Diaphorina citri are naturally infected with Wolbachia; therefore, investigating Wolbachia-mediated effects on D. citri fitness and CLas transmission required artificial reduction of this endosymbiont with the application of doxycycline. Doxycycline treatment of psyllids reduced Wolbachia infection by approximately 60% in both male and female D. citri. Psyllids treated with doxycycline exhibited higher CLas acquisition in both adults and nymphs as compared with negative controls. In addition, doxycycline-treated psyllids exhibited decreased fitness as measured by reduced egg and nymph production as well as adult emergence as compared with control lines without the doxycycline treatment. Our results indicate that Wolbachia benefits D. citri by improving fitness and potentially competes with CLas by interfering with phytopathogen acquisition. Targeted manipulation of endosymbionts in this phytopathogen vector may yield disease management tools. Graphical Abstract

Silencing of genes through RNA interference (RNAi) in insects has gained momentum during the past few years. RNAi has been used to cause insect mortality, inhibit insect growth, increase insecticide susceptibility, and prevent the development of insecticide resistance. We investigated the efficacy of topically applied dsRNA to induce RNAi for five Cytochrome P450 genes family 4 (CYP4) in Diaphorina citri. We previously reported that these CYP4 genes are associated with the development of insecticide resistance in D. citri. We targeted five CYP4 genes that share a consensus sequence with one dsRNA construct. Quantitative PCR confirmed suppressed expression of the five CYP4 genes as a result of dsRNA topically applied to the thoracic region of D. citri when compared to the expression levels in a control group. Western blot analysis indicated a reduced signal of cytochrome P450 proteins (45 kDa) in adult D. citri treated with the dsRNA. In addition, oxidase activity and insecticide resistance were reduced for D. citri treated with dsRNA that targeted specific CYP4 genes. Mortality was significantly higher in adults treated with dsRNA than in adults treated with water. Our results indicate that topically applied dsRNA can penetrate the cuticle of D. citri and induce RNAi. These results broaden the scope of RNAi as a mechanism to manage pests by targeting a broad range of genes. The results also support the application of RNAi as a viable tool to overcome insecticide resistance development in D. citri populations. However, further research is needed to develop grower-friendly delivery systems for the application of dsRNA under field conditions. Considering the high specificity of dsRNA, this tool can also be used for management of D. citri by targeting physiologically critical genes involved in growth and development.

Pear psylla, Cacopsylla pyricola (Förster) (Hemiptera: Psyllidae), is the most economically important pest of pears grown in Washington State. Standard conventional management programs involve season-long broad-spectrum insecticide sprays. Although the industry uses some tools that are not disruptive to biological control, such as kaolin clay and selective insecticides, they are additions to broad-spectrum insecticides instead of replacements. Conventional sprays suppress pear psylla through the spring and early summer; however, disruption of biological control leads to pear psylla outbreaks near harvest. In 2018 and 2019, we tested two season-long programs that used only selective approaches. The programs began with either kaolin clay or reflective plastic mulch and were followed by identical spray programs using only selective insecticides. Programs were compared with an industry standard conventional program that used numerous broad-spectrum insecticides throughout the season, and a check program with no insecticides for pear psylla. Experiments were conducted using replicated 40-tree plots in a research orchard near Wenatchee, WA with high pear psylla pressure. In both years, selective programs had similar pear psylla densities to the industry standard program and all had lower pear psylla densities and fruit injury than the check. Both selective programs had lower fruit injury than the industry standard in the first year, and similar injury to the industry standard in the second year. Our results suggest kaolin clay and reflective mulch can effectively suppress pear psylla populations and injury in the early season and support season-long selective management programs without the use of broad-spectrum insecticides.

Genome reduction in obligately intracellular bacteria is one of the most well-established patterns in the field of molecular evolution. In the extreme, many sap-feeding insects harbor nutritional symbionts with genomes that are so reduced that it is not clear how they perform basic cellular functions. For example, the primary symbiont of psyllids (Carsonella) maintains one of the smallest and most AT-rich bacterial genomes ever identified and has surprisingly lost many genes that are thought to be essential for its role in provisioning its host with amino acids. However, our understanding of this extreme case of genome reduction is limited, as genomic data for Carsonella are available from only a single host species, and little is known about the functional role of “secondary” bacterial symbionts in psyllids. To address these limitations, we analyzed complete Carsonella genomes from pairs of congeneric hosts in three divergent genera within the Psyllidae (Ctenarytaina, Heteropsylla, and Pachypsylla) as well as complete secondary symbiont genomes from two of these host species (Ctenarytaina eucalypti and Heteropsylla cubana). Although the Carsonella genomes are generally conserved in size, structure, and GC content and exhibit genome-wide signatures of purifying selection, we found that gene loss has remained active since the divergence of the host species and had a particularly large impact on the amino acid biosynthesis pathways that define the symbiotic role of Carsonella. In some cases, the presence of additional bacterial symbionts may compensate for gene loss in Carsonella, as functional gene content indicates a high degree of metabolic complementarity between co-occurring symbionts. The genomes of the secondary symbionts also show signatures of long-term evolution as vertically transmitted, intracellular bacteria, including more extensive genome reduction than typically observed in facultative symbionts. Therefore, a history of co-evolution with secondary bacterial symbionts can partially explain the ongoing genome reduction in Carsonella. However, the absence of these secondary symbionts in other host lineages indicates that the relationships are dynamic and that other mechanisms, such as changes in host diet or functional coordination with the host genome, must also be at play.

The spatial niche occupation of the Asian citrus psyllid, Diaphorina citri Kuwayama, 1908, was evaluated to determine its field colonization and food resource exploitation strategies in citrus groves. Mature grapefruit and sweet orange groves were surveyed as part of an area-wide program in 2009-2010 to determine D. citri population densities and between-tree distribution. In both cultivars, significantly more psyllids were found on perimeter trees throughout the study period suggesting a strong edge effect in D. citri distribution in the groves. D. citri densities and infestation levels gradually declined from the edge to the center of grove. Higher numbers of D. citri were recorded on trees located on the east and south sides of the groves than those on the west and north sides. Citrus groves located at the outer edge of the study with at least one side non-surrounded to other citrus groves harbored significantly more D. citri than groves located within the block cluster and entirely surrounded by other groves. In detailed field studies during 2012, infestation of D. citri started from border trees in the grove where possibly one generation is completed before inner trees become infested. In addition, psyllid densities decreased significantly with increasing distance from the grove edge. Using the selection index, D citri exhibited a strong niche occupation preference for border trees.

Ciriacremum is a genus of Ciriacreminae (Hemiptera, Sternorrhyncha, Psylloidea, Psyllidae) comprising 26 validly described species in the Afrotropical realm and only one in the Neotropics. Here we add four new species from Brazil, which are named and diagnosed, including illustrations and an identification key for adults: Ciriacremum lanceolatum sp. nov. (from Mato Grosso), C. maederae sp. nov. (from Amazonas), C. pollicigerum sp. nov. (from Amazonas) and C. roraima sp. nov. (from Roraima). Two other species are reported from Brazil (Amazonas and Pará) but not formally named due to insufficient material. Apart from C. setosum (from Nicaragua) and the Brazilian species treated in this paper, there are at least four additional undescribed species from the Neotropics (Colombia, Guyana, Panama, Suriname, and Venezuela), which are briefly discussed. Little is known about the immatures and host plants of the Neotropical species. The only confirmed host is Hymenaea sp. (Fabaceae) of C. maederae sp. nov. Adults were collected also on other fabaceous genera that are possible hosts. In the Neotropics, Ciriacremum is restricted to the tropics between Nicaragua in the north and Brazil (Mato Grosso) in the south.

The Asian citrus psyllid, Diaphorina citri Kuwayama, is the vector of the bacterial pathogen, Candidatus Liberibacter asiaticus, which is the causal agent of huanglongbing (HLB) in the United States. Both short-range and long-range dispersal of D. citri adults affect the spread of HLB; however, little is known about the long-range dispersal capabilities of D. citri in the field or the seasonality of flight behavior. In the present study, an in situ protein marking technique was used to determine the dispersal of D. citri by trapping marked adults under natural field conditions. D. citri movement from abandoned citrus groves to adjacent managed citrus groves was greatest during the spring and summer months and decreased significantly during the colder months (September–March). D. citri were able to traverse potential geographic barriers such as roads and fallow fields. In an experiment conducted to determine long-range dispersal capacity in the absence of severe weather events, D. citri were able to disperse at least 2 km within 12 d. Wind direction was not correlated with the number of marked psyllids captured, indicating substantial flight capability by D. citri. Finally, the number of marked psyllids captured increased with the density of emerging young leaves on surrounding trees. The results confirm that abandoned citrus groves in Florida serve as reservoirs for D. citri, which can disperse across long distances despite geographical barriers.

The development of insecticide resistance in Asian citrus psyllid, Diaphorina citri Kuwayama, populations is a serious threat to the citrus industry. As a contribution to a resistance management strategy, we developed a glass vial technique to monitor field populations of Asian citrus psyllid for insecticide resistance. Diagnostic concentrations needed to separate susceptible genotypes from resistant individuals were determined for cypermethrin (0.5 µg per vial), malathion (1.0 µg per vial), diazinon (1.0 µg per vial), carbaryl (1.0 µg per vial), carbofuran (0.1 µg per vial), methomyl (1.0 µg per vial), propoxur (1.0 µg per vial), endosulfan (1.0 µg per vial), imidacloprid (0.5 µg per vial), acetamiprid (5.0 µg per vial), chlorfenapyr (2.5 µg per vial), and fenpyroximate (2.5 µg per vial). In 2014, resistance to two carbamate insecticides (carbaryl and carbofuran), one organophosphate (malathion), one pyrethroid (cypermethrin), and one pyrazole (fenpyroximate) was detected in field populations of Asian citrus psyllid in Immokalee, FL. There was no resistance detected to diazinon, methomyl, propoxur, endosulfan, imidacloprid, and chlorfenapyr. The levels of insecticide resistance were variable and unstable, suggesting that resistance could be successfully managed. The results validate the use of the glass vial bioassay to monitor for resistance in Asian citrus psyllid populations and provide the basis for the development of a resistance management strategy designed to extend the efficacy of all classes of insecticides used for control of the Asian citrus psyllid.