Quarantine strategies for olive fruit fly (Diptera: Tephritidae): low-temperature storage, brine, and host relations

A dose-response relationship was not observed in olive fruit fly, Bactrocera oleae (Gmelin), larvae exposed to acetic acid concentrations (0-2.5%) used in commercial brine solutions to cure olives. Immersion in a 1% acetic acid brine solution impeded emergence of the immature stages. A 1-wk exposure of olives infested with olive fruit fly larvae to low-temperature storage as a postharvest treatment at 0-1 degree C resulted in 8% survival of the population, and exposures of 2 through 5 wk further reduced pupal and adult emergence to <1.0%. One- to 2-wk exposures at 2-3 degrees C resulted in a significant decrease in survival from 20 to 3%, respectively, and longer durations of 3-5 wk reduced survival to <1.0%. Mean daily fruit pulp temperatures in olives in the top, middle, and bottom of plastic bins stored at 2-3 degrees C decreased by 5-8 degrees C from the first to the second day. Lowest temperatures were observed in the top, and highest temperatures were observed in the middle layer of fruit, which attained a mean temperature of 3.8 degrees C on day 5. Laboratory choice tests showed that olive fruit fly oviposited at a higher rate in late season Mission olives that were green than in fruit that were in the red blush maturity stage in tests with 1- and 3-4-d exposure periods, and an increase in duration of exposure was related to an increase in the total number of ovipositional sites. Higher percentages of olive fruit fly third instars, pupae, and adults were reared from green fruit than from fruit in the red blush stage after a 1-d exposure to oviposition. Manzanillo olives were more attractive for oviposition by olive fruit fly than Mission olives, and significantly more third instars, pupae, and adults developed in Manzanillo fruit than in Mission fruit in the red blush stage. These differences were related to the better quality and higher flesh content of the Manzanillo versus Mission olives used in the tests.     

Psyttalia ponerophaga (Hymenoptera: Braconidae) as a potential biological control agent of olive fruit fly Bactrocera oleae (Diptera: Tephritidae) in California

The olive fruit fly, Bactrocera oleae (Rossi), is a newly invasive, significant threat to California's olive industry. As part of a classical biological control programme, Psyttalia ponerophaga (Silvestri) was imported to California from Pakistan and evaluated in quarantine. Biological parameters that would improve rearing and field-release protocols and permit comparisons to other olive fruit fly biological control agents were measured. Potential barriers to the successful establishment of P. ponerophaga, including the geographic origins of parasitoid and pest populations and constraints imposed by fruit size, were also evaluated as part of this investigation. Under insectary conditions, all larval stages except neonates were acceptable hosts. Provided a choice of host ages, the parasitoids' host-searching and oviposition preferences were a positive function of host age, with most offspring reared from hosts attacked as third instars. Immature developmental time was a negative function of tested temperatures, ranging from 25.5 to 12.4 days at 22 and 30 degrees C, respectively. Evaluation of adult longevity, at constant temperatures ranging from 15 to 34 degrees C, showed that P. ponerophaga had a broad tolerance of temperature, living from 3 to 34 days at 34 and 15 degrees C, respectively. Lifetime fecundity was 18.7 +/- 2.8 adult offspring per female, with most eggs deposited within 12 days after adult eclosion. Olive size affected parasitoid performance, with lower parasitism levels on hosts feeding in larger olives. The implications of these findings are discussed with respect to field manipulation and selection of parasitoid species for olive fruit fly biological control in California and worldwide.     

Olive fruit fly (Diptera: Tephritidae) populations in relation to region, trap type, season, and availability of fruit

Olive fruit fly, Bactrocera oleae (Gmelin), was monitored with adult captures by season and trap type, and was related to fruit volume and nonharvested fruit to elucidate the occurrence of the newly introduced pest in California. The highest numbers of adults captured in ChamP traps in olive trees, Olea europaea, were in October in an inland valley location, and in September in a coastal location. Comparisons of trap types showed that the number of olive fruit fly adults captured in Pherocon AM traps in a commercial orchard was significantly greater than in ChamP traps. A significantly greater number of females were captured in Pherocon AM traps with bait packets and pheromone lures than traps with pheromone lures alone, while a significantly greater number of adults and males were captured in traps with pheromone lures alone. Significantly more adults were captured in ChamP traps with bait packets and pheromone lures versus traps with bait packets alone. Fruit volume increased by four times from mid-June to mid-November. Olive fruit fly was found to oviposit on small olive fruit <1 cm3 shortly after fruit set, the maximum number of ovipositional sites per fruit occurred in October, and the greatest number of pupae and adults were reared from fruit collected in September and October. The highest numbers of pupae were collected from nonharvested fruit in March when high numbers of adults were captured in the same orchard.     

Insecticidal activity of strains of Bacillus thuringiensis on larvae and adults of Bactrocera oleae Gmelin (Dipt. Tephritidae).

The olive fly, Bactrocera oleae, is the key pest on olives in the Mediterranean area. The pest can destroy, in some cases, up to 70% of the olive production. Its control relies mainly on chemical treatments, sometimes applied by aircraft over vast areas, with their subsequent ecological and toxicological side effects. Bacillus thuringiensis is a spore-forming soil bacterium which produces a protein crystal toxic to some insects, including the orders of Lepidoptera, Diptera, and Coleoptera and other invertebrates. The aim of this study was to search for isolates toxic to B. oleae. Several hundred B. thuringiensis isolates were obtained from olive groves and olive presses in different areas of Greece, Sardinia (Italy), and Spain and from cooperating scientists throughout the world. Some isolates were found toxic only to adults or larvae and some to both stages of the olive fly. In addition, the most toxic isolates were assayed on Opius concolor Szepl. (Hym. Braconidae), the most important parasitoid of the olive fruit fly. Only 3 isolates out of 14 gave significant mortality against this parasitoid. Several of the most toxic crystalliferous isolates may contain novel toxins since they gave no PCR products when probed with primers specified for 39 known toxin genes.     

Psyttalia cf. concolor (Hymenoptera: Braconidae) for biological control of olive fruit fly (Diptera: Tephritidae) in California

The larval parasitoid, Psyttalia cf. concolor (Szépligeti), reared on Mediterranean fruit fly, Ceratitis capitata (Weidemann), by the USDA-APHIS-PPQ, Guatemala City, Guatemala, was imported into California for biological control of olive fruit fly, Bactrocera oleae (Gmelin), in olives, Olea europaea L. Mean percentage parasitism of olive fruit fly third instars infesting fruit in field cages ranged from 7.0 in Grapevine to 59.7 in Santa Barbara and in free releases ranged from 0 in Grapevine to 10.6 in Santa Barbara after 4- to 6-d exposures. In the laboratory, more parasitoids developed to adults in olive fruit fly larvae that were 11-13 d old than in larvae 8-10 d old. Adult parasitoids lived significantly longer when provided with water than adults without water in environmental chambers at 5 degrees C, 85% RH; 15 degrees C, 65% RH; 25 degrees C, 25% RH; and 35 degrees C, 25% RH. Adult parasitoids lived for 48 d with honey for food and water and 32 d with food and sugar solution at 15 degrees C and 65% RH. Survival of adult parasitoids without food and water in greenhouse tests was approximately 4 d in a simulated coastal climate and 1 d in a simulated inland valley climate and was significantly increased by providing food and water. The parasitoid did not develop in the beneficial seedhead fly, Chaetorellia succinea (Costa), in yellow star thistle. The rate of parasitism of walnut husk fly, Rhagoletis completa Cresson, larvae in green walnut husks was 28.4% in laboratory no-choice tests. In choice tests, the rate of parasitism of walnut husk fly versus olive fruit fly larvae in olives was 11.5 and 24.2%, respectively.     

Host plant selection and development in Spodoptera exigua: do mother and offspring know best?

We examined the ovipositional preference and larval development of Spodoptera exigua (Hübner) (Lepidoptera: Noctuidae) on two common hosts in southern California, Chenopodium murale L. (Chenopodiaceae) and Apium graveolens L. (Umbelliferae) to determine if female oviposition preference is correlated with offspring performance. Greenhouse oviposition choice tests indicated that S. exigua oviposit more frequently on C. murale than on A. graveolens. However under laboratory conditions, larvae reared on C. murale had longer development times, lower relative growth rate, and lower survivorship than larvae reared on A. graveolens. larval and pupal masses were significantly greater on A. graveolens than on C. murale. Furthermore, pupal masses were significantly greater for individuals reared on A. graveolens than on C. murale. Because pupal masses and adult fecundity are positively correlated for Spodoptera spp., the fitness of S. exigua on A. graveolens is likely to be substantially higher than its fitness on C. murale. Despite better larval performance on A. graveolens, previous results from choice tests with whole plants and leaf discs indicate that the highly mobile S. exigua larvae strongly prefer C. murale over A. graveolens. Hypotheses attempting to explain this lack of correlation between larval and adult host preference versus development and survival in this system are discussed.     

Microflora species and their volatile compounds affecting development of an alcohol dehydrogenase homozygous strain (Adh-I) of Bactrocera (Dacus) oleae (Diptera: Tephritidae)

Microflora species and volatiles emitted from artificial diets were examined from the larvae of three homozygous alcohol dehydrogenase (Adh) strains of the olive fruit fly, Bactrocera (Dacus) oleae (Gmelin), reared under identical conditions. Differences in volatile composition were detected when Adh-I homozygous larvae developed in a diet lacking the preservative p-hydroxybenzoic acid methyl ester (nipagin). Larval development of the Adh-I strain in the preservative-free diet was reduced by 50%, whereas pupal emergence was completely inhibited. The larval development and pupal emergence of Adh-F and Adh-S strains were not affected. Unique microorganisms with characteristic volatile profiles were isolated from the preservative-free diet of the Adh-I strain that were different from those, isolated from Adh-S, Adh-F, laboratory colony, and wild insect populations. Our results indicated that the variations in volatile composition of the artificial diets, and the inhibition of larval development and pupal emergence in Adh-I strain were related to changes in the microflora that developed in the diets of the Adh-I strain.     

Effect of larval density and host species on preimaginal development of Bactrocera tau (Diptera: Tephritidae)

The fruit fly, Bactrocera tau (Walker) (Diptera: Tephritidae), is an important pest of fruit and vegetable crops. In this study, host preference of B. tau females and the effects of host species and larval density on larval survival, pupal weight, adult emergence, and developmental duration were investigated on cucumber (Cucumis sativus L.), sponge gourd (Luffa cylindrical L. (Roem)), bitter gourd [Momordica charantia (Cucurbitaceae) L.], guava [Psidium guajava (Myrtaceae) L.], and tangerine [Citrus reticulata (Rutaceae) (Blanco)]. The results showed that females preferred to cucumber over other host species. Larval feeding experience affected subsequent host oviposition preference of adult females. Host species and initial larval density affected certain aspects of the biology of B. tau. Larval density negatively affected insect performance. Survival rates at low densities were significantly higher than that at high densities. Total developmental duration reduced at high larval densities. Cucumber was more suitable to larval growth. Larvae on cucumber grew faster and the puparia were heavier than that on other host species. Larval survival, pupation rate and adult emergence were higher on cucumber compared with those in other host species. Oviposition preference of adult females was correlated with performance of their offspring.     

Prevalence of Candidatus Erwinia dacicola in wild and laboratory olive fruit fly populations and across developmental stages

The microbiome of the olive fruit fly, Bactrocera oleae (Gmelin), a worldwide pest of olives (Olea europaea L.), has been examined for >100 yr as part of efforts to identify bacteria that are plant pathogens vectored by the fly or are beneficial endosymbionts essential for the fly's survival and thus targets for possible biological control. Because tephritid fruit flies feed on free-living bacteria in their environment, distinguishing between the transient, acquired bacteria of their diet and persistent, resident bacteria that are vertically transmitted endosymbionts is difficult. Several culture-dependent and -independent studies have identified a diversity of species in the olive fruit fly microbiome, but they have not distinguished the roles of the microbes. Candidatus Erwinia dacicola, has been proposed to be a coevolved endosymbiont of the olive fruit fly; however, this was based on limited samples from two Italian populations. Our study shows that C. Erwinia dacicola was present in all New and Old World populations and in the majority of individuals of all life stages sampled in 2 yr. Olive fruit flies reared on olives in the laboratory had frequencies of C. Erwinia dacicola similar to that of wild populations; however, flies reared on artificial diets containing antibiotics in the laboratory rarely had the endosymbiont. The relative abundance of C. Erwinia dacicola varied across development stages, being most abundant in ovipositing females and larvae. This uniform presence of C. Erwini dacicola suggests that it is a persistent, resident endosymbiont of the olive fruit fly.     

Population structure and colonization history of the olive fly, Bactrocera oleae (Diptera, Tephritidae)

The olive fly, Bactrocera oleae, is the major pest of olives in most commercial olive-growing regions worldwide. The species is abundant in the Mediterranean basin and has been introduced recently into California and Mexico, creating problems for quarantine protection and international trade. Here, we use nuclear microsatellite markers and mitochondrial sequences to examine the history of olive fly range expansion and colonization. Sampled populations span the current distribution of the olive fly worldwide, including South and Central Africa, Pakistan, Mediterranean Europe and Middle East, California, and Mexico. The Pakistani populations appear to be genetically well differentiated from the remaining populations, though rooting the origins of the species is problematic. Genetic similarity and assignment tests cluster the remaining populations into two genetic groups--Africa and a group including the Mediterranean basin and the American region. That Africa, and not the Mediterranean, is the origin of flies infesting cultivated olive is supported by the significantly greater genetic diversity at microsatellite loci in Africa relative to the Mediterranean area. The results also indicate that the recent invasion of olive flies in the American region most likely originated from the Mediterranean area.     

Evaluation of Fopius arisanus as a biological control agent for the olive fruit fly in California

1 The egg‐prepupal parasitoid Fopius arisanus (Hymenoptera: Braconidae) was evaluated in quarantine facilities as a potential biological control agent for the olive fruit fly Bactrocera oleae (Diptera: Tephritidae) in California, U.S.A. 2 Nontarget testing of two weed biological control agents confirmed that F. arisanus will not attack Tephritidae that feed in inflorescences or galls. It may, however, pose risks to native Tephritidae that feed in fruit. 3 Females preferentially oviposited in eggs, although first‐instar B. oleae were also attacked. Low lifetime reproductive potential and high rates of direct mortality inflicted on host eggs indicate that rearing on B. oleae may prove difficult. 4 In multiparasitized B. oleae, F. arisanus prevailed in competition against two species of larval–pupal parasitoids, Diachasmimorpha kraussii and Psyttalia concolor (both Hymenoptera: Braconidae). 5 The broad host‐range of F. arisanus with respect to fruit‐feeding Tephritidae may preclude its introduction to California, as may its low fecundity and its intrinsic competitive superiority over larva l–pupal parasitoids, which include specialists on B. oleae that are currently being introduced to California. High rates of direct mortality, however, point to potential uses in augmentative biological control. Whether or not F. arisanus is released in California, its biology as a parasitoid of B. oleae has been little studied to date and the results herein may be applied in other regions worldwide where B. oleae is a problem.     

幼虫密度对甜菜夜蛾生长发育与繁殖的影响

为了研究甜菜夜蛾Spodoptera exigua(Hübner)幼虫的密度对其发育及繁殖的影响,本实验观察了5种幼虫密度下（1,5,10,20,30头/瓶）,幼虫发育和成虫繁殖情况。结果表明：幼虫和蛹历期、存活率和蛹重均差异显著。幼虫和蛹历期均以20头/瓶的最短,1头/瓶的最长,其余随幼虫密度增加而延长;幼虫至蛹存活率以10头/瓶的最高,其余随幼虫密度增加而降低;1头/瓶的蛹最重,显著高于其他密度的,其余随幼虫密度增加而下降。尽管密度间成虫羽化率和产卵前期均无显著差异,但成虫产卵量、寿命和畸形率差异显著。1头/瓶的产卵量最多,其次为10头/瓶的,其余随幼虫密度增加而减少,30头/瓶的产卵量显著少于其他密度的; 密度在1～20头/瓶范围内,雌蛾寿命均较短,显著短于30头/瓶的,而雄蛾寿命以5头/瓶的最短,显著短于其他密度的（10头/瓶除外）,10头/瓶的次之,其余密度间差异不显著;不同幼虫密度下羽化的成虫畸形率差异显著,10头/瓶的最低,其余随幼虫密度增加而升高; 生命表结果表明甜菜夜蛾在10头/瓶下世代存活率和种群增长指数均最高,幼虫密度过低或过高均不利于种群增长;世代存活率（S）和种群增长指数（I）与幼虫密度之间的关系均呈抛物线关系：S =－0.2087x²+2.5694x+211.52 (R²=0.88),I=－0.0552x²+0.9166x+54.168 (R²=0.95)。结果提示幼虫密度影响甜菜夜蛾种群动态的重要生态因子之一。     

Mitotic and polytene chromosome analysis in Dacus oleae (Diptera: Tephritidae).

The present study constitutes the first attempt to construct a photographic map of the polytene chromosomes of Dacus oleae, a pest of the olive tree that causes serious financial damage in all olive oil producing countries. The map was constructed by using the larval fat body cells, the chromosomes of which are representative of the polytene chromosomes of other polytene tissues. In addition, the mitotic chromosomes of brain ganglia were examined, permitting tentative correlations between mitotic and polytene elements. This investigation shows that D. oleae is suitable for cytogenetic analysis in both mitotic and polytene chromosomes, a fact that may prove very useful for obtaining more detailed genetic information on the pest's natural populations.     

Female preference and larval performance of sunflower moth,Homoeosoma electellum,on sunflower pre‐breeding lines

The preference–performance hypothesis for insect herbivores predicts that adult females should preferentially choose hosts on which their offspring perform better. We tested this hypothesis for the sunflower moth, Homoeosoma electellum (Hulst) (Lepidoptera: Pyralidae), using 16 sunflower (pre‐breeding) lines, derived from a number of wild species of Helianthus, including Helianthus annuus L., Helianthus deserticola Heiser, Helianthus paradoxus Heiser, Helianthus praecox Engelm. & Gray ssp. hirtus (Heiser) Heiser, Helianthus praecox Engelm. & Gray ssp. runyonii (Heiser) Heiser, Helianthus petiolaris Nutt., Helianthus resinosus Small, and Helianthus tuberosus L. (Asteraceae), that are suitable for introducing wild sunflower germplasm into commercial cultivars. Female moths showed a range of ovipositional preference measures to the various lines. Combined data for three Helianthus species represented by multiple lines showed significant differences in female preference with respect to the parental species. Larval performance, determined by proportion of infested neonate larvae reaching the pupal stage, or mean pupal weight, varied across the lines and, as for the female preference data, also showed significant differences among the three parental Helianthus species represented by multiple lines. These data suggest that the characteristics in the pre‐breeding lines influencing female sunflower moth preference and larval performance likely originate from the parental species and may be consistently transferred to the derived pre‐breeding lines. Of particular note with regard to potential plant resistance mechanisms, lines derived from H. tuberosus showed consistent low preference–performance measures. Female preference and larval performance (for both measures) were strongly correlated, indicating that females preferred plants and lines on which larvae performed better, in support of the preference–performance hypothesis.     

Density and structure of Saissetia oleae (Hemiptera: Coccidae) populations on citrus and olives: relative importance of the two annual generations

Saissetia oleae (Olivier) (Hemiptera: Coccidae) populations were studied and compared in citrus (Citrus spp.) and olive (Olea europaea L.) groves to determine the number of generations, crawler emergence periods and changes in population density during the year. Ten citrus and four olive groves were sampled regularly between March 2003 and December 2005 in eastern Spain, covering an area of 10,000 km2. Each sample consisted of 16 branches and 64 leaves. Saissetia oleae populations presented a similar trend in both crops during the three years of study. Populations peaked in July, when crawlers emerged after the egg-laying period, and decreased during several months due to mortality of first instars in summer. A second crawler emergence period, with lower numbers and more variability from year to year, occurred between October and March. Populations did not increase during this period, probably because most eggs and crawlers perished during the winter and also because females that gave rise to this fall-winter generation were half as big and fecund as spring females. No differences were found between the size of mature females that had developed on citrus and on olives during the spring. Considering this population pattern, the best seasonal period to apply pesticides to control S. oleae would be at the end of July, when populations are synchronous, all crawlers have already emerged, and first instars predominate.     

Preference and performance linkage of a leaf-mining moth on different Salicaceae species

The relationship between oviposition prefer-ence and offspring performance of a leaf-mining moth (Paraleucoptera sinuella) on four Salicaceae species was investigated in 1997 and 1998. We observed the egg distribution pattern on different plant species in the field and carried out oviposition experiments in the laboratory to determine the preference of ovipositing females. We also examined larval survival, pupal mass, and developmental time to compare larval performance on each plant species. Egg density in the field differed significantly among plant species. However, egg density was not correlated exactly with demonstrated oviposition preference. No larvae could develop on two Salix species. This finding indicated that larval survival is the most critical index of larval performance. Larval performance on each plant species was correlated well with oviposition preference that was revealed by a no-choice experiment in the laboratory. However, this correlation was not found in the field. These results indicate that the preference–performance linkage that was observed under laboratory conditions, was not always maintained in the field. Received: September 25, 2000 / Accepted: April 27, 2001     

Oviposition preference,larval performance and adaptation of Trichoplusia ni on cabbage and cotton

Most female herbivores ensure to lay eggs where their offspring can develop successfully. The oviposition preferences of females affect strategies in pest management. In this study, the performance of two cohorts of Trichoplusia ni larvae on cabbage and cotton (after they had been transferred from their original host plants) were investigated. The preferences of female moth ovipositing and larval feeding on these two host plants were observed. The results indicated that plants significantly affected oviposition preference of the female adults and development and survival of larvae of T. ni. All females preferred to lay eggs on cabbage than cotton regardless from which host they originated. The detrimental effects of cotton on the development and survival of T. ni larvae originated from cabbage (CaTn) increased with the increase of the larval age when they were transferred. In addition, the host plant change did not significantly affect the development and survival of larvae of T. ni originating from cotton (CoTn). Larvae of CaTn preferred cabbage plants as compared to cotton plants, whereas larvae of CoTn did not show a significant choice. Although the adult females preferred laying eggs on cabbage, they did not show preferences between cotton and cabbage in a Y‐tube olfactometer test. The hypothesis of oviposition preference and performance of larvae was supported by the results of CaTn, whereas they not supported by those from CoTn. Based on these results, the strategy to manage this serious pest was discussed.     

Effects of host variety on blue willow beetle Phratora vulgatissima performance

Phratora vulgatissima adults exhibit feeding preferences among willow varieties, yet little is known of the effects of willow variety on larval and adult performance. The effects of host variety on adult fitness and fecundity as well as on larval mortality and development were studied under laboratory conditions for 35 willow varieties. The host variety significantly affected the number of days that adults survived and the rates of weight change. On average, males lost weight and females gained weight. The total number of eggs, eggs day¹, eggs clutch^?1 and the length of the oviposition period were also significantly affected by willow variety. Progeny from eggs laid by adults fed on the different willow varieties showed significant differences in days to pupation and pupal weight when subsequently reared on Salix × dasyclados or on the same variety as fed to the adults. However, there was no correlation between these parameters on S. × dasyclados and the variety fed to adults. The willow variety fed to larvae significantly affected larval mortality (four varieties caused 100% mortality), the shape of larval growth curves (as measured by predicted final weight and time to half the final weight), the number of days to pupation and pupal weight. There were significant positive correlations between previously determined adult P. vulgatissima feeding preferences of the 35 willow varieties and the following: number of eggs laid, length of the oviposition period, larval mortality and development and change in adult weight. There was a considerable degree of variation in these correlations and some varieties did not follow the general trend indicated by the size or sign of particular correlations, for example, having a high feeding preference ranking yet few eggs laid, low larval weight and longer time to pupation. The differences found between varieties for adult and larval performance in conjunction with previously established feeding preferences offer great potential for utilising plant resistance to P. vulgatissima as a means of strategic control. Despite the general correlation of feeding preference and performance parameters, the results found here suggest that it appears to be possible for plant breeders to circumvent this trend.