Mating Behavior and Performance in the Two Cosmopolitan Mealybug Species Planococcus citri and Pseudococcus calceolariae (Hemiptera: Pseudococcidae)

Mating strategies of mealybugs were investigated using two heterogeneric cosmopolitan species as case study: Planococcus citri (Risso) and Pseudococcus calceolariae (Maskell). Male mating behavior of the studied species differed in respect to mate selection, and frequency and duration of copulation. Females played an active role in mate selection, by either facilitating or resisting copulation. This is a first evidence of female sexual active behavior in scale insects. In both species, male predisposition to mate was affected by previous exposition to light. The frequency of courtship and copulation decreased with the age of females for Pl. citri but not for Ps. calceolariae. Mating frequency increased with female colony density in both species.     

Molecular and Morphological Identification of Mealybug Species (Hemiptera: Pseudococcidae) in Brazilian Vineyards

Mealybugs (Hemiptera: Pseudococcidae) are pests constraining the international trade of Brazilian table grapes. They damage grapes by transmitting viruses and toxins, causing defoliation, chlorosis, and vigor losses and favoring the development of sooty mold. Difficulties in mealybug identification remain an obstacle to the adequate management of these pests. In this study, our primary aim was to identify the principal mealybug species infesting the major table grape-producing regions in Brazil, by morphological and molecular characterization. Our secondary aim was to develop a rapid identification kit based on species-specific Polymerase Chain Reactions, to facilitate the routine identification of the most common pest species. We surveyed 40 sites infested with mealybugs and identified 17 species: Dysmicoccus brevipes (Cockerell), Dysmicoccus sylvarum Williams and Granara de Willink, Dysmicoccus texensis (Tinsley), Ferrisia cristinae Kaydan and Gullan, Ferrisia meridionalis Williams, Ferrisia terani Williams and Granara de Willink, Phenacoccus baccharidis Williams, Phenacoccus parvus Morrison, Phenacoccus solenopsis Tinsley, Planococcus citri (Risso), Pseudococcus viburni (Signoret), Pseudococcus cryptus Hempel, four taxa closely related each of to Pseudococcus viburni, Pseudococcus sociabilis Hambleton, Pseudococcus maritimus (Ehrhorn) and Pseudococcus meridionalis Prado, and one specimen from the genus Pseudococcus Westwood. The PCR method developed effectively identified five mealybug species of economic interest on grape in Brazil: D. brevipes, Pl. citri, Ps. viburni, Ph. solenopsis and Planococcus ficus (Signoret). Nevertheless, it is not possible to assure that this procedure is reliable for taxa that have not been sampled already and might be very closely related to the target species.     

Predation of the apefly,Spalgis epius (Lepidoptera: Lycaenidae) on citrus mealybug,Planococcus citri (Hemiptera: Pseudococcidae)

The citrus mealybug, Planococcus citri (Risso) (Hemiptera: Pseudococcidae) is a serious pest of economically important crops worldwide. The apefly, Spalgis epius (Westwood) (Lepidoptera: Lycaenidae) is a potential predator of various species of mealybugs. Earlier investigation on its daily preying capacity and preference for prey stages on P. citri is incomplete. Hence, a study was conducted to find out the daily prey consumption ability and preference for prey stages by different larval instars of S. epius reared on P. citri in the laboratory. Through the 8-day developmental period with four larval instars, the daily prey consumption of S. epius increased from the first to the seventh day and decreased on the eighth day prior to the prepupal stage. Generally, there was a significant difference in the prey consumption on different days. When the prey stages were offered separately, the first to fourth instar larva of S. epius consumed, respectively, a mean of 199.6, 722.6, 1908.8, and 4625.6 eggs or 21.5, 77.0, 168.5, and 670.5 nymphs or 3.2, 7.2, 16.0, and 35.1 adults of P. citri. When an S. epius larva was fed on P. citri eggs, nymphs and adults separately, it consumed a mean of 7456.7 eggs, 937.6 nymphs, or 62.3 adults during its entire development. When the prey stages were offered all together, a single S. epius larva consumed 2618.4 eggs, 170.4 nymphs, and 39.7 adults of P. citri throughout its entire development. The study revealed that S. epius is a voracious predator of P. citri and thus could be utilized as a major biological control agent.     

Pheromone-based mating disruption of Planococcus ficus (Hemiptera: Pseudococcidae) in California vineyards

Experiments were conducted to test a mating disruption program for the mealybug Planococcus ficus (Signoret) (Hemiptera: Pseudococcidae) in California vineyards. The sprayable, microencapsulated formulation of the racemic sex pheromone lavandulyl senecioate was applied with an air-blast sprayer, using three and four applications in 2003 and 2004, respectively. Mating disruption was combined with an application of buprofezin (2004) in June. Compared with a no-pheromone control, there were significantly lower season-long trap catches of adult males, season-long mealybug densities (2003 only), and crop damage in mating disruption plots. The amount of mealybug reduction and mechanisms that resulted in lower crop damage in mating disruption plots is discussed. In samples taken during the growing season (April to September), mealybug density was only 12.0 +/- 15.6 and 31.1 +/- 11.6% lower in the mating disruption plots than in control plots in 2003 and 2004, respectively. In the mating disruption treatment, mealybug egg production was significantly lower (2003 only), as were the proportion of ovisacs and crawlers produced. There was no treatment impact on percentage of parasitism. Mealybug density influenced treatment impact. In 2004, vines were categorized as having low, medium, or high mealybug densities during a preapplication survey. After treatment application, mealybug density was reduced by 86.3 +/- 6.3% on vines in the low mealybug density category, but it was unchanged on vines in the high density category. Another factor that reduced treatment impact was the relatively short effective lifetime of the sprayable formulation.     

Identification of Planococcus citri (Homoptera: Pseudococcidae) and the description of a new species

Planococcus pacificus sp.n. is described from the South Pacific, Australia and South East Asia. P.citricus is synonymized with P.citri. A key and illustrations are given to separate the species discussed from each other and from the similar P.ficus group.     

Genetic Characterization of a Putative Densovirus from the Mealybug Planococcus citri

Total genomic DNA preparations from the citrus mealybug, Planococcus citri, contained a DNA band corresponding to 5.5 kilobases. This DNA was a linear molecule and was cloned into pUC18. Nucleotide sequence determination indicated that it was the replicative form of a densovirus, most closely related to the virus from Periplaneta fuliginosa (smokybrown cockroach). Received: 25 April 2001 / Accepted: 3 May 2001     

Baseline susceptibility of Planococcus ficus (Hemiptera: Pseudococcidae) from California to select insecticides

Between 2006 and 2008, 20 populations of Planococcus ficus (Signoret), from Coachella and San Joaquin Valleys of California were measured in the laboratory for susceptibility to buprofezin, chlorpyrifos, dimethoate, methomyl, and imidacloprid. Toxicity was assessed using a petri dish bioassay technique for contact insecticides and by a systemic uptake technique for imidacloprid. Mixed life stages were tested for susceptibility to all insecticides except for buprofezin, which was measured against early and late instars (first, second, and third). Dose-response regression lines from the mortality data established LC50 and LC99 values by both techniques. Responses of populations from the two geographical locations to all five insecticides varied, in some cases significantly. Variations in susceptibility to each insecticide among sample sites showed a sevenfold difference for buprofezin, 11-fold to chlorpyrifos, ninefold to dimethoate, 24-fold to methomyl, and 8.5-fold to imidacloprid. In spite of susceptibility differences between populations, baseline toxicity data revealed that all five insecticides were quite effective based on low LC50s. Chlorpyrifos was the most toxic compound to Planococcus ficus populations as shown by lowest LC50s. Buprofezin was toxic to all immature stages but was more potent to first instars. The highest LC99 estimated by probit analysis of the bioassay data of all 20 populations for each compound was selected as a candidate discriminating dose for use in future resistance monitoring efforts. Establishment of baseline data and development of resistance monitoring tools such as bioassay methods and discriminating doses are essential elements of a sustainable management program for Planococcus ficus.     

Effect of Host Plants on Honeydew Production of an Invasive Mealybug, Phenacoccus solenopsis (Hemiptera: Pseudococcidae)

Honeydew production plays a key role in mutualism between the mealybugs and ants. However, no studies have focused on the amount and circadian rules of honeydew excreted by Phenacoccus solenopsis Tinsley, a new invasive species which has conditional mutualism with Solenopsis invicta Buren in China. To address this problem, we measured the weight and estimated honeydew production in all stages of development of the invasive mealybug, P. solenopsis, as well as its honeydew production on tomato (Solanum lycopersicun), Hibiscus rosa-sinensis, and cotton (Gossypium sp.) for 24 h. The honeydew excreted by each instar of the mealybug in H. rosa-sinensis was measured for 2 weeks. Our results revealed that the weight of mealybugs significantly varied at different development stages. Host plants had no significant effect on the weight of nymphs, although the weight of a single adult reared on S. lycopersicun was significantly heavier than those reared on H. rosa-sinensis and G. sp. The amount of honeydew excreted by the 1st instar nymphs in S. lycopersicum was significantly greater than that on H. rosa-sinensis and G. sp. Each instar mealybug produced more honeydew when fed with S. lycopersicum compared with H. rosa-sinensis and G. sp. The amount of honeydew excreted by mealybugs when provisioned with H. rosa-sinensis was no different from mealybugs provisioned with G. spp. while in the same instar. The amount of honeydew excreted by the 1st and 2nd instar nymphs was not significantly different on the same host plant. However, there was a significant difference between the 3rd instar nymph and the adult. The amount of honeydew excreted by a single adult when provisioned with H. rosa-sinensis decreased from 3085.3 μg to 572.0 μg in 2 weeks. The 2nd instar nymph, 3rd instar nymph, and adult excreted honeydew more frequently during the day than at night, while the frequency of honeydew excretion of the 1st instar nymph had no significant difference between daytime and night.     

Occurrence of Planococcus minor Maskell (Hemiptera: Pseudococcidae) in cotton fields of northeast region of Brazil

The mealybug Planococcus minor Maskell is a significant pest of more than 250 cultivated plants belonging to at least 80 families. Among the host plants several Malvaceae species are attacked, including the genera Gossypium. In 2005 and 2006 the mealybug P. minor was observed infesting cotton fields in the arid area of northeast region of Brazil. The occurrence was registered in high densities and in several farms resulting in mortality of plants in some cases. This species is not cited as a cotton pest in any region of Brazil where cotton grows, although it is enlisted in the Global Pest and Disease Database. Hence, there is a lack of studies what makes this pest of great deal to control.     

Potential of South African entomopathogenic nematodes (Heterorhabditidae and Steinernematidae) for control of the citrus mealybug, Planococcus citri (Pseudococcidae)

Planococcus citri, the citrus mealybug, is the most important species of mealybug known to infest citrus in South Africa. Various laboratory bioassays were conducted to determine the potential of entomopathogenic nematodes to control P. citri. Adult female P. citri were screened for susceptibility to six indigenous nematode species. P. citri was found to be most susceptible to Steinernema yirgalemense and Heterorhabditis zealandica, causing 97% and 91% mortality, respectively. The development of nematodes after infecting adult female P. citri showed both H. zealandica and S. yirgalemense were able to complete their life cycles inside the host. Further bioassays illustrated a linear relationship between mealybug mortality and the concentration of nematodes applied, with the highest level of control using a concentration of 80 infective juveniles (IJs)/insect. As nematodes would be used as an above-ground application to control P. citri in citrus orchards, available water is a major limiting factor. Insecticidal activity proved to be dependent on the available surface moisture after nematode application. The water activity (a(w)) bioassay indicated that S. yirgalemense to be two times more tolerant to lower levels of free water, with a(w)(50)=0.96 and a(w)(90)=0.99, compared to H. zealandica with a(w)(50)=0.98 and a(w)90=1.0. After application, nematodes have a limited time frame in which to locate and infect hosts, as the level of available free water gradually decreases, as trees dry out. S. yirgalemense proved able to locate and infect P. citri quicker than H. zealandica. Nematode activity was not significantly affected when exposed to 15°C, 20°C and 25°C. IJs were able to infect P. citri at an exposure time as short as half an hour. Results also showed that the first 2-4h post application is the most decisive time for establishing successful infection of mealybugs. This is the first report on the potential use of nematodes for the control of P. citri.     

The passionvine mealybug,Planococcus minor (Maskell) (Hemiptera: Pseudococcidae), and its natural enemies in the cocoa agroecosystem in Trinidad

Planococcus minor (Maskell) is native to South Asia, but it is also present in several Neotropical locations including the island of Trinidad in the southern Caribbean. The mealybug poses a serious threat to uninfested countries in this region as well as the mainland U.S. As part of an effort to gather much needed information on P. minor, 33 cocoa (Theobroma cacao L.) field sites on the island were surveyed in 2006 with a view to assess the occurrence and pest status of the mealybug. P. minor was identified from 20 field sites, indicating that it was well distributed across the island on this crop, which appeared to be a reliable indicator host plant. Infestation levels were generally low and populations were sparsely distributed across the field sites categorized into three habitat types. The following year, nine field sites were surveyed for natural enemies of P. minor using laboratory-infested potatoes in sentinel traps. Species from four insect orders and six families were collected and identified. The major predators belonged to the families Cecidomyiidae and Coccinellidae. Two primary parasitoids, Leptomastix dactylopii Howard (Encyrtidae) and Coccidoxenoides perminutus (Girault) (=Pauridia peregrina Timberlake, =Coccidoxenoides peregrinus (Timberlake)) (Encyrtidae), were reared from different mealybug stages, along with several hyperparasitoids. The primary parasitoids were probably introduced fortuitously. These diverse natural enemies were recovered throughout the sampling period from the different habitat types. The identification of key natural enemies associated with P. minor has important implications for the implementation of biological control in newly infested areas.     

Endomitosis in the mealy bug,Planococcus citri (Homoptera: Coccoidea)

Endomitosis in the Malpighian tubules of the mealy bug Planococcus citri (Risso) is described. The stages are identified on the basis of the length of the chromosomes and the distance between the sister chromatids or chromosomes. The appearance of the chromosomes in the various stages of endomitosis is compared to that in other hemipteran insects. During anaphase and telophase of endomitosis the ends of the sister chromatids and chromosomes tend to stay together longer than the other parts. It is suggested that in holokinetic chromosomes special regions for holding the chromatids together are concentrated near the ends of the chromosomes.Supported by grant GB1585 from the National Science Foundation, Washington, D.C.     

Resistance of Citrus and Related Genera to Diaphorina citri Kuwayama (Hemiptera: Liviidae)

The present study was developed to evaluate the resistance of the following genotypes of Citrus and related genera to this pest: ‘Pera,’ ‘Natal’, and ‘Washington Navel’ oranges (Citrus sinensis), ‘Marsh Seedless’ grapefruit (Citrus paradisi), hardy orange ‘Rubidoux’ (Poncirus trifoliata), kumquat (Fortunella margarita Swingle), citrumelo ‘Swingle’ (C. paradisi x P. trifoliata), and citrange ‘Troyer’ (P. trifoliata x C. sinensis). The experiments were performed in greenhouses with plants grafted onto ‘Rangpur’ lime (Citrus limonia) and placed individually in voile cages. The preference for oviposition in a no-choice test, and the effect of genotype were evaluated. The egg-adult cycle was monitored to determine the effect of genotype on the biology of the insect. Poncirus ‘Rubidoux’ was the least preferred genotype for oviposition; reduced number of eggs was also found to occur on citrange ‘Troyer’, and ‘Marsh Seedless’ was the genotype with the most eggs. No significant variation in the duration of the embryonic period was observed; however, a difference in the viability of eggs was found, with the lowest egg viabilities on ‘Swingle.’ Kumquat and ‘Marsh Seedless’ genotypes were correlated with increased durations of the nymphal phase, however, there was no difference in the survival of this phase. Fecundity of females on ‘Troyer’, ‘Swingle’, and kumquat was reduced. Considering all of the evaluated parameters, it was concluded that cultivars of sweet orange are the most susceptible genotypes to Diaphorina citri. Regarding oviposition, P. trifoliata ‘Rubidoux’ showed resistance of the antixenosis type.     

The Genetic Structure of an Invasive Pest,the Asian Citrus Psyllid Diaphorina citri (Hemiptera: Liviidae)

The Asian citrus psyllid Diaphorina citri is currently the major threat to the citrus industry as it is the vector of Candidatus Liberibacter, the causal agent of huanglongbing disease (HLB). D. citri is native to Asia and now colonizes the Americas. Although it has been known in some countries for a long time, invasion routes remain undetermined. There are no efficient control methods for the HLB despite the intensive management tools currently in use. We investigated the genetic variability and structure of populations of D. citri to aid in the decision making processes toward sustainable management of this species/disease. We employed different methods to quantify and compare the genetic diversity and structure of D. citri populations among 36 localities in Brazil, using an almost complete sequence of the cytochrome oxidase I (COI) gene. Our analyses led to the identification of two geographically and genetically structured groups. The indices of molecular diversity pointed to a recent population expansion, and we discuss the role of multiple invasion events in this scenario. We also argue that such genetic diversity and population structure may have implications for the best management strategies to be adopted for controlling this psyllid and/or the disease it vectors in Brazil.     

Reproductive biology of Pseudococcus maritimus (Hemiptera: Pseudococcidae)

This study characterized several parameters associated with reproductive behaviors of a Californian population of the grape mealybug, Pseudococcus maritimus (Ehrhorn). Experiments with caged adults confirmed that females from this population could not reproduce parthenogenetically. Both sexes were capable of mating multiple times on the same day and on sequential days. Median times between copulations were short (< 10 min) on the first day that males were presented with females, but tended to increase with sequential copulation events. Unmated females lived for up to 19 wk, whereas mating and oviposition resulted in reduced longevity. We hypothesized that constant exposure to pheromone might stimulate males to leave their cocoons sooner and increase their activity levels compared with males that were not exposed to pheromone, resulting in shortened life-spans. However, whereas exposure to pheromone did stimulate males to emerge slightly sooner than control males, pheromone exposure had no effect on the longevity of males. The implications of the results of these experiments for pheromone-based methods of detection and control of mealybugs are discussed.     

Nonreplication of heterochromatic chromosomes in a mealy bug,Planococcus citri (Coccoidea: Homoptera)

In males of mealy bugs with the lecanoid chromosome system, the paternal set of chromosomes becomes heterochromatic in early embryogeny. In males of the mealy bug, Planococcus citri, the heterochromatic (H) set in testis sheath cells and in most of the oenocytes apparently did not replicate while the euchromatic (E) set was undergoing several cycles of endoreplication. In third instar males, testis sheath cells in endoanaphase and endotelophase exhibited 5H and either 40 or 80E chromosomes. The increase in the number of E chromosomes was attributed to the replication of only the E chromosomes. Oenocytes of third instar males had 0, 5, or 10H chromosomes and from 10 to 240E chromosomes. The oenocytes with 5H chromosomes had a mean of 50.8E chromosomes, and those with 10H chromosomes had a mean of 155.6E chromosomes. Nuclear and cell fusion was considered as a means of producing the various numbers of H and E chromosomes in oenocytes, and it was concluded that although nuclear fusion probably took place, the differences between the number of H and E chromosomes was at least in part due to replication of only the E chromosomes. The size of the H chromosomes was about the same in all the testis sheath cells and the oenocytes irrespective of the level of endopolyploidy for the E set. These H chromosomes apparently did not increase in polyteny, because they were only about half the size of the H chromosomes in prophase I of spermatogenesis. The significance of the nonreplication of the H set and the control of nonreplication are briefly discussed.This study was aided by a grant (GB-1585) from the National Science Foundation, Washington, D.C.